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ZOOTAXA

2101

Taxonomic revision and of the -feeding (Brullé, 1832) group (Coleoptera: Chrysomelidae: : ) and analysis of their potential in biological control of Tamarisk

JAMES L. TRACY & THOMAS O. ROBBINS

Magnolia Press Auckland, New Zealand JAMES L. TRACY & THOMAS O. ROBBINS Taxonomic revision and biogeography of the Tamarix-feeding Diorhabda elongata (Brullé, 1832) species group (Coleoptera: Chrysomelidae: Galerucinae: Galerucini) and analysis of their potential in biologi- cal control of Tamarisk (Zootaxa 2101) 152 pp.; 30 cm. 11 May 2009 ISBN 978-1-86977-359-5 (paperback) ISBN 978-1-86977-360-1 (Online edition)

FIRST PUBLISHED IN 2009 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2009 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use.

ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

2 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Zootaxa 2101: 1–152 (2009) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2009 · Magnolia Press ISSN 1175-5334 (online edition)

Taxonomic revision and biogeography of the Tamarix-feeding Diorhabda elongata (Brullé, 1832) species group (Coleoptera: Chrysomelidae: Galerucinae: Galerucini) and analysis of their potential in biological control of Tamarisk

JAMES L. TRACY1 & THOMAS O. ROBBINS 1USDA-ARS, , Soil and Water Research Laboratory2808 East Blackland Road, Temple, TX 76502. E-mail: [email protected], [email protected]

Dedication

This work is dedicated to Culver Jack DeLoach, Jr., our long time friend and mentor in weed biological control research who has inspired us with his dedication and energy for bringing the natural marvel of biological control of invasive into the process of native ecosystem restoration, especially in his last two decades of research and implementation in the use of Diorhabda as tamarisk biological control agents in North America.

Table of contents

Abstract ...... 4 Резюме ...... 4 Introduction ...... 4 Materials and methods ...... 9 ...... 15 Diorhabda Weise, 1883 ...... 15 Diorhabda elongata-Group...... 18 Key to the Sexes of the Diorhabda elongata Group: Adult External Characters ...... 39 Key to the Species of the Diorhabda elongata Group: Adult Males ...... 39 Key to the Species of the Diorhabda elongata Group: Adult Females ...... 40 Diorhabda elongata (Brullé, 1832) ...... 44 Diorhabda carinata (Faldermann, 1837) ...... 57 Diorhabda sublineata (Lucas, 1849) REVISED STATUS ...... 74 Diorhabda carinulata (Desbrochers, 1870) ...... 86 Diorhabda meridionalis Berti & Rapilly, 1973 NEW STATUS ...... 102 Implications Regarding Biological Control of Tamarisk ...... 128 Opportunities for Further Research...... 134 Acknowledgements ...... 135 Literature cited ...... 136

Accepted by G. Morse: 4 Mar. 2009; published: 11 May 2009 3 Abstract

The primarily Palearctic Diorhabda elongata species group is established for five Tamarix-feeding sibling species (tamarisk ): D. elongata (Brullé, 1832), D. carinata (Faldermann, 1837), D. sublineata (Lucas, 1849) REVISED STATUS, D. carinulata (Desbrochers, 1870), and D. meridionalis Berti & Rapilly, 1973 NEW STATUS. Diorhabda koltzei ab. basicornis Laboissière, 1935 and D. e. deserticola Chen, 1961 are synonymized under D. carinulata NEW SYNONYMY. Illustrated keys utilize genitalia, including male endophallic sclerites and female vaginal palpi and internal sternite VIII. Distribution, comparative biogeography, biology, and potential in biological control of Tamarix in North America are reviewed. Diorhabda elongata is circummediterranean, favoring Mediterranean and temperate forests of Italy to western . Diorhabda carinata resides in warm temperate , deserts, and forests of southern Ukraine south to Iraq and east to western . Diorhabda sublineata occupies Mediterranean from France to North and subtropical deserts east to Iraq. Diorhabda carinulata primarily inhabits cold temperate deserts of Mongolia and China west to Russia and south to montane grasslands and warm deserts in southern . Diorhabda meridionalis primarily occupies maritime subtropical deserts of southern and Iran to Syria. Northern climatypes of D. carinulata are effective in Tamarix biological control, especially in the Great Basin desert. Diorhabda elongata is probably best suited to Mediterranean woodlands of northern California. Northern climatypes of D. carinata may be best suited for central U.S. grasslands. Diorhabda sublineata, D. meridionalis, and southern climatypes of D. carinata and D. carinulata may each be uniquely suited to areas of the southwestern U.S.

Key words: Diorhabda elongata species group; Chrysomelidae; Taxonomy; Comparative Biogeography; Biology; Host Range; Tamarix; Tamarisk; Saltcedar; Weed Biological Control; Sibling Species; Hybrid Morphology; Morphometry; Genitalic Phenogram; Biomic Dendrogram; Habitat Suitability Index Models

Резюме

Видовая группа Diorhabda elongata основана для пяти палеарктических, питающихся на Tamarix, видов- двойников (тамарисковые жуки): D. elongata (Brulle, 1832), D. carinata (Faldermann, 1837), D. sublineata (Lucas, 1849) REVISED STATUS, D. carinulata (Desbrochers, 1836) и D. meridionalis Berti & Rapilly, 1973 NEW STATUS. Diorhabda koltzei ab. basicornis Laboissiere, 1935 и D. e. deserticola Chen, 1961 синонимизированы с D. carinulata NEW SYNOMYMY. Иллюстрированный определитель использует гениталии, включая мужской эндофалус, женские вагинальные пальпы и внутренний стернит VIII. Рассмотрено распространение, сравнительная биогеография, биология и возможности использования для биологического контроля Tamarix в Северной Америке. Diorhabda elongata является циркум-средиземноморским видом, предпочитающим средиземноморские и умеренные леса Италии до западной Турции. Diorhabda carinata обитает в теплых умеренных остепненных биотопах, пустынях и лесах южной Украины и далее на юг до Ирана и восток до Западного Китая. Diorhabda sublineata обитает в лесах Средиземноморья от Франции до Северной Африки и субтропических пустынях на восток до Ирака. Diorhabda carinulata в основном населяет холодные и умеренные пустыни Монголии и Китая и далее на запад до России и на юг до горных степей и теплых пустынь южного Ирана. Diorhabda meridionalis населяет приморские субтропические пустыни южного Пакистана и Ирана до Сирии. Северный климатип D. carinulata является эффективным агентом биологического контроля Tamarix, особенно в пустынях Большого Бассейна. Diorhabda elongata, вероятно, наиболее подходит для контроля Tamarix в средиземноморских лесах Северной Калифорнии. Северный климатип D. carinata может быть наиболее подходящим для травянистых биоценозов центральной части США. Diorhabda sublineata, D. meridionalis, южный климатип D. carinata и D. carinulata могут быть подходящими для различных мест юго-запада США.

Introduction

Tamarix-feeding beetles in the genus Diorhabda Weise (1883), or tamarisk beetles, are probably the most damaging specialized defoliators of Old World tamarisks (Kulinich 1962; Sinadsky 1968; Tomov 1979; Samedov and Mirzoeva 1985; Tian et al. 1988; Bao 1989; Sha and Yibulayin 1993; Myartseva 1999; Mityaev and Jashenko 1999, 2007; DeLoach et al. 2003b). Consequently, tamarisk beetles are valued as current and

4 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS potential biological control agents for invasive tamarisks in North America (DeLoach et al. 2003b, 2004; Lewis et al. 2003b, Milbrath and DeLoach 2006a; Carruthers et al. 2006, 2008). Accurate taxonomic recognition of the species of tamarisk beetles and their associated biogeographic characteristics could be crucial to their successful utilization in biological control of tamarisk across the widely varying biogeography of the western North American tamarisk invasion. Many taxonomists consider the tamarisk beetles to comprise a single species, D. elongata (Brullé, 1832), with three to four subspecies ranging from and southwestern to Mongolia and western China (Wilcox 1971, Riley et al. 2003, Bieńkowski 2004, Lopatin et al. 2004). However, Berti and Rapilly (1973) removed two tamarisk beetles, D. carinata (Faldermann, 1837) and D. carinulata (Desbrochers, 1870), from synonymy with one another, and by implication with D. elongata. Their findings were based primarily upon diagnostic differences in the male endophalli (eversible internal sacs of the aedeagi) in the holotypes of D. carinata and D. carinulata, but they did not characterize the endophallus of D. elongata. We characterize the endophallus of D. elongata and corroborate the findings of Berti and Rapilly (1973). We also find several additional diagnostic male and female genitalic characters by which to distinguish these species, remove D. sublineata (Lucas, 1849) from synonymy with D. elongata, and elevate D. carinulata meridionalis Berti and Rapilly (1973) to species status as D. meridionalis Berti and Rapilly. A complex of five fully diagnosable species of tamarisk beetles is established as forming the Diorhabda elongata species group: D. elongata, D. carinata, D. sublineata, D. carinulata and D. meridionalis. Four species of tamarisk beetles (excluding D. meridionalis) are recently introduced for biological control of tamarisk in the . We provide the first detailed distributional data for all five species of tamarisk beetles, revealing their unique biogeographic characteristics and providing a basis for further ecogeographic studies. New data on the biogeography of tamarisk beetles are used in matching each species to different regions of the North American tamarisk invasion using hand-fitted habitat suitability index models. The Eurasian tamarisks T. ramosissima Ledebour, T. chinensis Loureiro and their hybrids (mixed populations of pure lines and hybrids all referred to as T. ramosissima/T. chinensis) are highly invasive in arid and semi-arid riparian areas of the western U.S. (Gaskin and Schaal 2002, 2003), where they are the second most dominant woody riparian after native cottonwoods (Populus deltoides Barton ex Marshall) (Friedman et al. 2005). de Candolle is invasive in riparian areas of California (DiTomaso 1998, Gaskin and Schaal 2003). Populations of T. canariensis Willdenow/T. gallica Linnaeus are minor invasives along the Gulf Coast of and Louisiana, and they commonly hybridize with T. ramosissima/T. chinensis in Texas (Gaskin and Schaal 2002, 2003; John Gaskin, USDA/ARS, Sydney, MT, pers. comm.). (Linnaeus) Karsten (athel tamarisk) is an evergreen ornamental (DeLoach 1990) planted primarily in subtropical areas of the southwestern U.S. and northern where it commonly escapes vegetatively and sometimes propagates by and is showing the potential to become more invasive (Walker et al. 2006) as it is along the Finke River in Australia (Griffen et al. 1989). Naturalized hybrids of T. aphylla and T. ramosissima/T. chinensis have recently been found in the southwestern U.S. which could possibly contribute to the invasive potential of Tamarix (Gaskin and Shafroth 2005). Dominance of exotic tamarisks is facilitated by their pre-adaptation to low soil moisture riparian habitats that have widely increased due to stream flow regulation practices (Glen and Nagler 2005, Lite and Stromberg 2005, Stromberg et al. 2007). Tamarisks also invade unregulated river, spring, and marsh systems where they displace a variety of woody and herbaceous riparian communities, and can alter ecosystem function to the detriment of native biota, including endangered species (DiTomaso 1998, Lovich and DeGouvenain 1998, Tracy and DeLoach 1999, DeLoach et al. 2000, Stenquist 2000, Dudley and DeLoach 2004, Kennedy et al. 2005, Birken and Cooper 2006, Whiteman 2006, Whitcraft et al. 2007). Invasive tamarisk could cost an estimated US $7–16 billion in lost ecosystem function in the U.S. from 2000–2055, mainly as a result of losses in hydropower (on ) and wildlife habitat (Zavaleta 2000). Great effort and expense is being directed towards control of tamarisk to both reduce its use of water and help restore native riparian ecosystems (Hart 2003, Shafroth et al. 2005, Carruthers et al. 2008). Herbicidal treatment of individual tamarisk is often practiced (Duncan 2003), especially in southern California and

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 5 Arizona (Barrows 1993, Brock 2003). Large-scale control of tamarisk by aerial herbicidal application, sometimes in combination with mechanical removal or prescribed burning, is the prevalent control method in New Mexico and west Texas (Hughes 2003, McDaniel and Taylor 2003, Hart et al. 2005). Over 100 species of Old World tamarisk specialist herbivores that attack T. ramosissima in Asia (Kovalev 1995) are absent in North America, and the lack of these herbivores is an often overlooked factor contributing to the invasiveness of tamarisk and its competition against comparatively herbivore-stressed native flora (DeLoach et al. 2000, Dudley et al. 2000). The objective of classical biological control of tamarisk is to re-associate tamarisk in North America with its Old World Tamarix-specific natural enemies in order to reduce tamarisk populations below thresholds requiring other control methods. Tamarisk biological control can contribute to native ecosystem restoration by permanently suppressing tamarisk without collateral damage to native riparian flora, with low cost, and without need of re-application (DeLoach 1990, 1997; Gould and DeLoach 2002). The use of tamarisk beetles in biological control was recently initiated, and they are already filling a critical herbivore niche in North American riparian ecosystems by continually suppressing tamarisk over large areas in Nevada, Utah and Wyoming (DeLoach et al. 2004; DeLoach and Carruthers 2004a; Carruthers et al. 2006, 2008; Dave Kazmer, USDA/ARS, Sydney, MT, pers. comm.), allowing increase in understory , while the tamarisk beetles serve as forage for insectivorous (Longland and Dudley 2008) and small mammals (Dudley 2005a). Populations of tamarisk beetles from China and Kazakhstan that we studied for biological control were initially identified as D. elongata deserticola Chen (1961) or “D. elongata” (DeLoach et al. 2003b; Lewis et al. 2003a, 2003b; Milbrath and DeLoach 2006a; Milbrath et al. 2007; DeLoach et al. in prep.). These identifications were obtained from several chrysomelid taxonomists, including Igor Lopatin (Belarus State University, Minsk, Belarus), Alexander Konstantinov (USDA-ARS SEL, Beltsville, MD), Shawn Clark (Brigham Young University, Provo, Utah), Steven Lingafelter (USDA-ARS SEL, Beltsville, MD), and Richard White (USDA-ARS SEL, retired). Permits for release of D. elongata in North America were first issued by USDA- and Plant Health Inspections Service (APHIS) in 1999 (USDA-APHIS 1999, DeLoach et al. 2000). In this taxonomic revision, we synonymize D. e. deserticola under D. carinulata, the northern tamarisk . Populations of northern tamarisk beetles from 44°N in northwestern China (Fukang and Turpan) and near Shelek (= Chilik), Kazakhstan, were released (as D. elongata) into North America for biological control of tamarisk in 2001. Diorhabda carinulata has established well on T. ramosissima/T. chinensis at sites north of 37°N in Nevada, Utah, Colorado, and Wyoming, defoliating large areas of tamarisk, especially in Nevada, Utah (DeLoach et al. 2004; Carruthers et al. 2006, 2008), western Colorado (Dan Bean, Colorado Department of Agricutlure, Grand Junction, CO, pers. comm.), and Wyoming (D. Kazmer, pers. comm.). However, these populations of D. carinulata exhibit asynchrony of daylength induced diapause with seasonal changes at southern sites in New Mexico and Texas, where they initially failed to establish (DeLoach et al. 2004, Bean et al. 2007a). Populations of tamarisk beetles that we studied from , , and Uzbekistan were originally identified as D. elongata by taxonomists A. Konstantinov and/or I. Lopatin. They all have diapause characteristics more adaptive to the southern U.S. (Bean and Keller in prep), they are all host specific to tamarisk (Milbrath and DeLoach 2006a, 2006b; Herr et al. 2006, in prep.), and they have all been released (as D. elongata) in North America with permits issued by USDA-APHIS for each specific population source. In this taxonomic revision, we confirm that populations of D. elongata from near Sfakaki (Crete) and Posidi Beach, Greece are the true D. elongata, the Mediterranean tamarisk beetle. Diorhabda elongata was established in 2004 on T. ramosissima/T. chinensis in west Texas and T. parviflora in northern California. But the Mediterranean tamarisk beetle has established poorly or not at all where it was released on T. ramosissima/ T. chinensis in eastern New Mexico and many locations in west and north Texas from 2004–2007. The population of “D. elongata” we studied from near , Tunisia is here identified as D. sublineata, the subtropical tamarisk beetle. Diorhabda sublineata was first released onto T. canariensis/T. gallica in south Texas in 2005 (Patrick Moran, USDA/ARS, Weslaco, TX, pers. comm.) but it has not yet established. Further releases of D. sublineata from near Marith, Tunisia are planned on T. chinensis/T. canariensis in south and

6 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS west Texas in 2009. The population of “D. elongata” we studied from near Qarshi (= Karshi), Uzbekistan is here classified as D. carinata, the larger tamarisk beetle. Diorhabda carinata was first released on T. ramosissima/T. chinensis in north Texas in 2006 (Jerry Michels, Texas AgriLIFE Research, Bushland, TX, pers. comm.) and appeared to be establishing near Seymour, Texas in 2008 (Charles Randal, USDA/APHIS, Olney, TX, pers. comm.). A population of “D. elongata” studied from near Buxoro (=Buchara), Uzbekistan (Herr et al. in prep.) was later found to be a mixture of D. carinata and D. carinulata. Diorhabda meridionalis, the southern tamarisk beetle, is primarily found in extreme southern and southwestern Iran, and it is yet to be studied for its potential in biological control. The list below summarizes the correlation between the revised taxonomy for tamarisk beetles and “D. elongata” population/ecotype designations used in a number of reports and publications by us and others in relation to tamarisk biological control (see recent citations under “D. elongata“ in the synonymies for each Diorhabda species account).

Scientific name Proposed common name “D. elongata”, “saltcedar ” population/ecotype designation D. carinulata northern tamarisk beetle Fukang and Turpan, China; Chilik, Kazakhstan; Buchara, Uzbekistan (part); D. e. deserticola, also “tamarisk leaf beetle” D. elongata Mediterranean tamarisk beetle Crete (nr Sfakaki) and Posidi Beach, Greece D. sublineata subtropical tamarisk beetle Sfax, Tunisia and southeast of Marith, Tunisia D. carinata larger tamarisk beetle Qarshi, Uzbekistan; Buchara, Uzbekistan (part) D. meridionalis southern tamarisk beetle south Iran (not yet introduced into USA)

Releases of tamarisk beetles in southern California, Arizona, and along the in western New Mexico, are currently delayed until concerns can be resolved regarding safety of tamarisk biological control to nesting habitats of the federally endangered southwestern , Empidonax traillii Audubon subspecies extimus Phillips, which will nest in tamarisk (see DeLoach et al. 2000, Dudley and DeLoach 2004). Efforts are continuing to establish tamarisk beetles across widely varying biogeographic areas invaded by tamarisk in the western U.S, but apparent adaptation problems are still being encountered in many southern areas. A basic understanding of the native biogeography of all the species of tamarisk beetles would be valuable in identifying their potential to adapt to different regions. Ongoing taxonomic research can be essential to the success of biological control programs. Precise species identifications are necessary to properly characterize genetic variability in biological control agents regarding: (1) risks to non-target vegetation; (2) efficacy and host preferences among target hosts; and (3) biogeographic adaptations at a variety of spatial scales (see also Schauff 1992). A recent example of the importance of ongoing taxonomic research in biological control is the discovery that a common biological control agent of exotic thistles in North America and Australia, the thistle rosette , Trichosirocalus horridus (Panzer) (Coleoptera: Curculionidae: Ceutorynchinae), actually comprises a complex of three sibling species, each apparently differing in the thistle species preferred as hosts (Alonso-Zarazaga and Sánchez-Ruiz 2002). An inability to taxonomically distinguish similar species of candidate biological control agents has led to lengthy delays in successful implementation of biological control programs. Such delays result from not recognizing and utilizing interspecific variability in the efficacy of potential agents (Gordh and Beardsley 1999). A more accurate appreciation of the interspecific variability in tamarisk beetles, especially in regard to their biogeographic characteristics, should facilitate more timely and effective utililization of their desirable traits in biological control of tamarisk. Our primary objectives are to clarify the number of morphologically diagnosable species of tamarisk beetles, and to stabilize their taxonomic status. To this end, we review all currently recognized specific and subspecific taxa of tamarisk-feeding Diorhabda. Sibling, or cryptic, species are difficult or impossible to distinguish based solely on external morphology. But, sibling species are often distinctly different in genitalic

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 7 morphology, distribution, behavior, ecology, and genetics, which is evidence of their reproductive isolation in nature. Our revision examines data on genitalic morphology, ecology, behavior and distribution, which are often the most reliable means of distinguishing sibling species, even with current genetic methods (Bull et al. 2006). The foundation of our revision is the geographic distribution of morphological character states of the genitalia, especially the endophallus, which are useful in distinguishing sibling species. Berti and Rapilly (1973) were among the first to use the gross morphology of the endophallus in distinguishing sibling species of Chrysomelidae in their restoration of the species D. carinata and D. carinulata from synonymy with D. elongata. Additional examples of taxonomic and morphological studies involving the endophallus in Chrysomelidae are given by Mann and Crowson (1996) and (1997). In the last 16 years, chrysomelid taxonomists have found the morphology of individual sclerites of the endophallus useful in distinguishing members of several sibling species complexes, including the nymphaeae (Linnaeus) species group (Galerucinae) (Lohse 1989), the Oulema melanopus (Linnaeus) species group (Criocerinae) (Berti 1989, Siede 1991, Hansen 1994), the flavipes Fabricius species group () (Duhaldeborde 1999), and the Cryptocephalus marginellus Olivier species group (Sassi 2001). In addition to characterizing the endophallic sclerites of tamarisk beetles, we also characterize structures of the female genitalia that are useful in separating similar species in Galerucinae, such as internal sternite VIII (e.g., LeSage 1986), spermathecae (e.g., LeSage 1986, Biondi and D’Alessandro 2003), and vaginal palpi (e.g., Baselga and Novoa 2005). Inaccurate identification of sibling species creates great confusion in the biological and ecological data published for these species. A current example of such confusion is seen in recent studies of sibling leaf beetle species in the species group. Lohse (1989) used a combination of five morphological characters (including endophallic sclerites) in distinguishing four sympatric members of the G. nymphaeae species group in central Europe (Warchalowski 2003, Bieńkowski 2004). Application of Lohse’s (1989) taxonomic characters is needed to clarify the identity of a number of these species in several biological and ecological investigations, including “G. sagittariae (Gyllenhal)” in Finland (Hippa and Koponen 1986; Nokkala and Nokkala 1994, 1998; Nokkala et al. 1998; but see Kangas 1991) (possibly G. k er s te n si Lohse and other species), “G. nymphaeae” host races from Polygonaceae in the Netherlands (Pappers et al. 2001, 2002a, 2002b; Pappers and Ouborg 2002) (possibly Lohse’s [1989] two Polygonum and Rumex host races of G. aquatica [Geoffroy]), and “G. nymphaeae” in North America (Otto and Wallace 1989, Nokkala et al. 1998, Cronin et al. 1999) (possibly one or more new species; the type locality of G. nymphaeae is Sweden according to Silfverberg [1974]). Similar taxonomic confusion over the identity of sibling species has sometimes clouded research in biological control programs (Schauff 1992, Clarke and Walter 1995, Schauff and LaSalle 1998). Another objective of this revision is to avoid further taxonomic confusion in future biological studies of D. elongata and its sibling species and to clarify the identity of these species as far as possible in the previous literature. We provide illustrated keys to sexes and species of adults of the D. elongata species complex and a genitalic phenogram showing morphological groupings based on similarity in key genitalic characters. Male genitalic hybrid morphologies are characterized for progeny from laboratory crossings of D. sublineata × D. elongata (F1 hybrids) and their F1 hybrids (F2 hybrids). Extensive annotated bibliographies are included under the synonymies for each species account that clarify numerous misapplications of names in the literature for more than 100 years. Distribution maps display (Olson and Dinerstein 2002), point localities for selected Tamarix spp., and point localities of Diorhabda species derived from examined specimens and literature records for which we could assign identifications based on species distribution patterns. Descriptive statistics are presented for elevation, latitude, distance to ocean (continentality) and percent distribution across biomes for field collection data of each Diorhabda species. We analyze similarities in biomes inhabited among the Diorhabda species and selected Tamarix species invasive in North America, providing biomic dendrograms and three dimensional biomic principal coordinate analysis scatter plots. Descriptive statistics for biogeographic variables are used to construct biomic dendrograms and hand-fitted habitat suitability index models to examine the comparative biogeography of the D. elongata group. We

8 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS provide geocoordinates used in the distribution maps that are essential to future development of species distribution models using climatic data. Label data from the literature and examined specimens are given. Information on the distribution, taxonomy, host plants, ecology, phenology, behavior, development, reproduction, biogeography, and potential in tamarisk biological control is reviewed for each tamarisk beetle species. New distribution and host records are reported. Finally, we summarize the implications of our research regarding biological control of tamarisk.

Materials and methods

Morphology. Genitalia are dissected (abbreviated “diss.” in material examined) from mounted specimens after first soaking the specimens in warm water for several hours to soften them before removal of the body from the pin or point. In males, the abdomen is usually removed from the body prior to extraction of the genitalia under a binocular stereoscope. Dissections are done in water over a yellow clay substrate to aid in positioning of the genitalia. Following removal of a male’s abdomen, the aedeagus is first pulled from the opening at the base of the abdomen to allow access to the endophallus (Fig. 14; END). We were unable to inflate the endophalli with a needle and pipette using a method similar to that of Silfverberg (1974). Occasionally, we everted the uninflated endophallus from the median lobe (ML) into its natural position by gently teasing the endophallus out through the distal dorsal ostium (DO, or apical orifice) of the aedeagus (Fig. 14) with a hooked minuten pin (inserted into a match stick). However, the eversion procedure is very tedious and is often unsuccessful, resulting in tearing of the endophallic membrane and sclerites, especially in older specimens. We usually employed the faster technique of dissecting out the endophallic sclerites in a non-everted (inverted) endophallus using the following procedure. An insect pin is used to first make a longitudinal slit along the entire length of the ventral membrane (VM) of the median lobe of the aedeagus (Fig. 14). The pin is then inserted through this slit at the basal foramen (BF, or basal orifice) and used to push the distal tip of the inverted endophallus out from the median lobe. Fine forceps are used to gently pull the inverted endophallus completely out of the median lobe, and the base of the inverted endophallus is carefully torn free from the membrane at the apex of the median lobe. The removed inverted endophallus is splayed open to reveal the endophallic sclerites. A hooked minuten pin is used to longitudinally slice the endophallic membrane from the torn opening at its base. Before slicing the endophallus, it is oriented such that, while the base is pointed downwards, the palmate (dorsal) endophallic sclerite (PES) is at the left and the elongate (ventral) endophallic sclerite (EES) is on the right in order to avoid damage to any linear connecting (lateral) endophallic sclerite (CES) (Figs. 14, 16). We develop terminology for endophallic sclerites of the D. elongata group (Figs. 14–33), and the preceding general terminology for endophallic sclerites in parentheses follows that of Mann and Crowson (1996). Extraction of the female genitalia (Figs. 34–38) normally involves first removing the abdomen. The last abdominal tergite and the internal sternite VIII (IS VIII) are then severed from connections to other abdominal tergites and sternites and the digestive tract to allow their removal with the attached oviduct, vaginal palpi (VP), and spermatheca (SP) (Fig. 34). Occasionally the spermatheca becomes detached in the abdomen after removal of the oviduct and has to be retrieved separately with forceps. In living specimens, the stalk of internal sternite VIII can sometimes be seen as a dark Y-shaped area through the transluscent last visible abdominal sternite (Lewis et al. 2003b, Fig. 1G). Also, in living specimens, the vaginal palpi and the apical lobe of internal sternite VIII can be seen between the last visible abdominal sternite and tergite after they are spread apart with forceps. Terminology of female genitalia (Figs. 34–43) follows that of LeSage (1986) and Konstantinov (1998). Duckett (2003, Fig. 5) illustrates the orientations of internal sternite VIII, vaginal palpi (as gonacoxae) and spermatheca with respect to one another in another galerucine beetle, Pedilia sirena Duckett. Camera lucida drawings are made at 80x magnification for endophallic sclerites of males and internal sternite VIII, spermathecae and vaginal palpi of females. Following examination of dissected genitalia, they

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 9 are stored in glycerin in polyethylene genitalia vials (4 mm × 10 mm) that are placed on the pin beneath the specimens. Removed abdomens are glued to points beneath the body of the specimens. Digital photographs are made of the habitus of adults with the aid of a stereomicroscope. Dissected specimens are assigned unique numbers, and label data, species identifications, and geocoordinates are entered into a database. Morphometrics. Only adults identified through examination of genitalia are used in measurements. Specimens measured are representative of the entire geographic range of each species, with from 15 to 61 individuals measured for each external character (Table 2) and from 12 to 29 individuals measured for each genitalic character (Tables 3 and 4). A calibrated ocular micrometer is used in making measurements of adults at 12.8x and 32x magnification and genitalia at 80x magnification. The blade or ridge of the elongate endophallic sclerite (EES) is measured as the total length of raised ridge (Figs. 14–28; LB). The extent of the blade is not as clearly distinguishable from the lateral view of the EES in D. elongata (Fig. 19, LB) as compared to other species of the group (Figs. 20–23). Most of the blade in D. elongata may be demarcated by a spindle-shaped outline (Fig. 24—Methoni,) seen from dorsal view of the EES, but the apex of the blade may be more fully demarcated by a single median dark line seen from either a certain dorso-lateral view (Fig. 24—Methoni, DLV) or the dorsal view (Fig. 24—nr ). The length of the spined (or toothed) area of the blade along the EES is measured as the distance from the basal tip of the sclerite to the most apical spine (Figs. 14–28; SL). If the EES is armed with a single distal spine (as is often the case for D. elongata), the length of the spined area is assigned a value of 0.01 mm. Accurate measurement of the blade length and spined area length of the EES can be important in distinguishing laboratory produced D. sublineata/D. elongata hybrids. We test for significant differences in some general external and key genitalic morphometric variables, especially looking for any morphometric discontinuities between the species. Various measurements of the female genitalia, including the vaginal palpi (VP) and internal sternite VIII (IS VIII), are illustrated in Figures 34–38. The frequency distributions of a few morphometric variables (e.g., width of elytra at widest point) fail the test for normality. For consistency, all data are analyzed using the nonparametric Two-Way Kruskal-Wallis Test on ranks using ANOVA and Fisher’s Protected LSD test to separate means of ranks (SAS Institute 2005). Scatter plots for measurements of the elongate endophallic sclerite are made with the Microsoft Excel Spreadsheet software. Stenophenetic analysis. Methods of cluster analysis used in numerical taxonomy (Sneath and Sokal 1973) are employed in analyzing phenetic similarities among the D. elongata species group. In contrast to the standard approach in phenetic analyses in which a large number of characters (ca. 60) is selected from throughout the body of each taxonomic unit for phenogram construction, we select only characters that are informative in species diagnosis, in this case eight characters from the male and female genitalia, to construct genitalic phenograms. We narrow the characters selected for phenetic analysis in a manner similar to that used in cladistic analyses in which only characters considered to be phylogenetically informative are used (Pankhurst 1991). We adopt the new term stenophenetic analysis to distinguish this approach from typical broader phenetic analysis. A rectangular data matrix of genitalic character state profiles is constructed with qualitative interval states for the eight key genitalic characters standardized to range from zero to one. An average taxonomic dissimilarity matrix, based on the average taxonomic distance coefficient, and a Pearson product-moment correlation similarity matrix, based on the Pearson product-moment correlation coefficient, are computed from the data matrix (NTSYSpc Interval Data [SIMINT] module [Rohlf 2006]) (see Sneath and Sokal [1973] for details on calculation of coefficients). Both the dissimilarity and similarity matrices are used in separate cluster analyses for the species with three standard clustering algorithms: unweighted pair-group method using arithmetic averages (UPGMA), complete-link method, and single-link method (NTSYSpc SAHN module). Genitalic dissimilarity and similarity phenograms are produced from each cluster analysis

(NTSYSpc plot module). The cophenetic correlation coefficient, rcoph, is calculated as a measure of goodness of fit for each phenogram to the original dissimilarity or similarity matrix from which it was derived through a particular method of cluster analysis (rcoph values from 0.80–0.90 indicate a good fit for the cluster analysis) (NTSYSpc Cophenetic values and Matrix Comparison Plot [MxComp] modules).

10 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Distribution. Cities and other precise geographic features given on locality labels of examined specimens or in literature records are assigned geocoordinates and updated place names using databases of the U.S. National Geospatial-Intelligence Agency (NGA) and the U.S. Board on Geographic Names accessed from internet geoname servers such as the GEOnet Names Server (NGA 2006). Locality label data giving distances and directions from landmarks and elevations are assigned geocoordinates with the aid of global elevation grid (1 km grid resolution), stream and road map layers, and distance measuring utilities using ArcMap software. Point location data are plotted in a Miller cylindrical projection for publication using ArcMap. Precision of point geocoordinates derived from precise locality label or literature data are in the range of ca. ± 5 km. Point geocoordinates derived from general locality data (precision much less than ca. ± 5 km) are noted as approximate locations. Geocoordinates and updated place names for label data of examined specimens are given in brackets under the Material Examined section for each species. Specimens that we examined with previously published identifications are noted in the Material Examined and Distribution sections of each species. Specimens from collection localities with previously published identifications that we did not examine are listed with geocoordinates and their probable identities are discussed under the heading Distribution-Unconfirmed Records for each species. Biogeography. Descriptive. The distribution of each species of Diorhabda is mapped and described in relation to biogeographic realms, biomes, , elevation, latitude, distance to the ocean (continentality), and the distributions of a few selected known and potential hosts of the genus Tamarix (comprised of ca. 90 species; Zhang and Zhang 1990, Qiner and Gaskin 2006). Realms, biomes and ecoregions are mapped with the ecoregions Arcview shapefile map of Olson and Dinerstein (2002). Approximate elevations in meters for plotted point locations are obtained from the GLOBE 1 km base digital elevation model ArcMap grid (Hastings et al. 1999). Distances to ocean are measured in kilometers with ArcMap software. Old World point distribution data for Tamarix spp. are obtained from a variety of regional and local floras (e.g., Post and Dinsmore 1932; Corti 1942; Rusanov 1949; Schiman-Czeika 1964; Baum 1967b, 1978, 1983; Qaiser and Ghafoor 1979; Baum and Townsend 1980; Qaiser 1983; De Martis et al. 1984, 1986; Browicz 1991; Boratyński et al. 1992; Zieliński 1994), ecological literature (e.g., Danin 1981, Hoberlandt 1981, Izco et al. 1984, Salinas et al. 2000, Omar et al. 2002, Kurschner 2004) and our herbarium records from Kazakhstan and western China (DeLoach unpublished). Data from published “dot” distribution maps are scanned, georeferenced according to their projection, and the geocoordinates queried using ArcMap software. For example, maps of Baum (1978) and Zieliński (1994) are in the Miller Cylindrical projection while those of Rusanov (1949) and Boratyński et al. (1992) are in the Bonne projection. Most North American locality data on T. ramosissima/T. chinensis are from Friedman et al. (2005). Additional North American Tamarix spp. locality data are primarily from online hebarium databases (e.g., Calflora, Seinet, and Tropicos) and our own herbarium specimens. Mexican locality data are less certain regarding both species identifications and possible ornamental status of T. ramosissima/T. chinensis (possibly confused with T. canariensis/T. gallica)at some locations, and data are included from De León González and Vásquez Aldape (1991). Primary habitats for each Diorhabda spp. are identified based upon frequency of collections, relative abundance with regard to sibling species, and reports of damage to Tamarix. Areas with possibly distinct climatypes (climatic ecotypes; Turesson 1925) that have potential use in biological control of tamarisk are identified and discussed in detail under the heading Potential in Tamarisk Biological Control for each species and in the concluding section, Implications Regarding Biological Control of Tamarisk (see for further explanation). Our assessments of the suitability of various biomes/elevation/latitude/distance to ocean combinations for Diorhabda spp. should be viewed with caution because the evenness of sampling effort across differing habitats and for differing species with presence-only distribution data are not controlled. Additional analyses are in progress with climatic data to produce presence-only species distribution models. Biomic Analysis. Methods of cluster analysis and principal coordinate analysis (PCoA) used in numerical ecology (Legendre and Legendre 1998) are employed in analyzing similarities in biomic occurrence state profiles (biomic profiles) among species of the D. elongata group and Tamarix spp. invasive in North

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 11 America. Rectangular data matrices of biomic profiles are constructed with semiquantitative states standardized to range from zero to one that indicate the degree of indigenous occurrence in various biomes: 0 - no record; 0.5 - single record (minor presence); 1 - multiple records (major presence). In view of the lack of systematic surveys for Diorhabda and Tamarix species in each , we consider more than one locality record as representing potentially more than a minor presence within a biome. We also calculate the biomic specialization index (BSI) (Hernández Fernández and Vrba 2005), which is the number of biomes inhabited by a given species, a low BSI indicating the highest biomic specialization. We modify BSI to represent the sum of biomic occurrence states for each species and use biomes as classified by Olson and Dinerstein (2002). From biomic profiles for Diorhabda species and Diorhabda and Tamarix species, we calculate biomic dissimilarity matrices and dendrograms with associated rcoph values using NTSYSpc (Rohlf 2006) in a manner similar to that described above for construction of the genitalic phenograms. However, the data matrices of biomic profiles are first normalized with the transformation log(x +1) (NTSYSpc Transformation module) and biomic Bray-Curtis dissimilarity matrices are then constructed using the Bray-Curtis (percentage difference) distance coefficient (NTSYSpc Interval Data [Simint] module), which eliminates double zero values from being counted as similarities in habitats (for details on calculation of Bray-Curtis coefficient, see distance coefficient D14 in Legendre and Legendre [1998]). The biomic Bray-Curtis dissimilarity matrix and the genitalic average taxonomic dissimilarity matrix (discussed previously) for the five Diorhabda species are analyzed for a positive correlation using a one-tailed Mantel test in which the significance level of the normalized Mantel statistic, rM, is determined using a Monte Carlo randomization procedure with 9,999 ≥ permutations (NTSYSpc, Matrix Comparison Plot [MxComp] module; test for probability random rM observed rM, including observed rM with random values in test). PCoA analysis is used to visualize biomic relationships among Diorhabda and Tamarix species. First, eigenvectors and eigenvalues are computed (NTSYSpc Eigen module) from the double centered biomic Bray- Curtis dissimilarity matrix (NTSYSpc Dcenter module). (A lack of negative PCoA axis eigenvalues did not necessitate using the square-root transformation for values in the Bray-Curtis dissimilarity matrix.) Eigenvalues of species along the first three PCoA eigenvector axes are displayed in a three dimensional biomic PCoA scatter plot (NTSYSpc Mod3D module). Loadings of the biomes on the first six PCoA axes are computed a posteriori using Spearman rank-order coefficients (Proc Corr; SAS Institute 2005) for ranks of the species in biomes versus ranks of the species in PCoA eigenvector axes. Habitat Suitability Index Models. Habitat suitability index (HSI) models estimate the habitat suitability of geographic areas for a given species based upon the subjectively or objectively appraised value of habitat indicator variables (suitability indices). HSI models were originally developed in the 1980’s to assess habitat suitability for various wildlife species by USDI Fish and Wildlife Service (1980, 1981) and were later coupled with GIS at a landscape level (e.g., Larson et al. 2003). We develop hand-fitted HSI models (5 minute grid resolution) to depict a first rough approximation of the relatively most suitable tamarisk beetle species for a given area. Four variables or biogeographic descriptors that we consider as encompassing intangible life requisites related to bioclimatic suitability for each Diorhabda species are employed in our HSI models: biome, latitude, elevation and distance to the ocean (continentality). Variable parameters employed in the model are from descriptive statistics of field collection data for each Diorhabda species assembled in this revision (locality data from specimens dissected in 2008 are not included in the models). ArcMap, ArcInfo and the above mentioned elevation and biome grids are used to create ArcGIS grids for each of the four HSI model variables and Diorhabda field collection data at 5 minute (0.08333 degree) grid cell resolution to facilitate computer processing and to approximate the level of precision in the less precise Diorhabda locality data. ArcInfo is used to calculate a grid of distance to ocean (continentality) using an Euclidean distance function between 5 minute resolution ocean grid cells (including the and Mediterranean Sea) and land grid cells derived from the above mentioned 1 km resolution elevation grid. A program macro for ArcMap is made to calculate a latitude grid using a point shapefile made from elevation land grid cells. Distribution points of Diorhabda that fall into 5 minute grid cells of incorrect values (e.g.,

12 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS ocean grid cells or incorrect biome grid cells) in the generalized biome and elevation grids are moved to adjacent grid cells of the appropriate value. The adjusted distribution points of each Diorhabda species are then converted into 5 minute Diorhabda locality grids used to query or sample values from the 5 minute grids of each of the four variables using ArcInfo. The resulting tables of 5 minute Diorhabda grid cell locations and grid cell values for each variable are used in calculating descriptive statistics in the form of schematic box plots for continuous variables (Proc Boxplot [Boxstyle = Schematic]; SAS Institute 2005) and frequency percentages for distribution of each Diorhabda species across eight biomes (Proc Freq; SAS Institute 2005; biome frequency percentage equals the number collections of a Diorhabda species within a biome divided by total numbers of collections for the species across all biomes). Descriptive statistics used in calculating habitat suitability indices are plotted using the Microsoft Excel Spreadsheet software. Lack of absence data and lack of even sampling across ranges of variables in native habitats precludes detailed statistical comparisons of means or frequencies. From descriptive statistics for the above four 5-minute variable grids, a total of four suitability index

(SI1–SI4) grids and one biomic relative suitability index (SI5) grid are calculated for each species. The biomic relative suitability index calculates the percent presence in a biome relative to data for other Diorhabda species in the biome and provides extra weighting for the biome variable in the model. A final HSI value is 1/n calculated as the geometric mean ([X1 × X2 ×… × Xn] ) of these five grids, giving a final HSI value of zero if any single SI variable is zero (Larson et al. 2003). Modeled values of the five suitability indices for each species are plotted against corresponding environmental variables using Microsoft Excel Spreadsheet software and the conditional statements and equations used in Arc Macro Language of ArcInfo for calculating HSI model grids are given below:

Suitability index (SI) grids Calculation per 5 minute grid cell for each Diorhabda speciesa latitudinal suitability index if within species interquartile range for latitude, then = 1; else if between max/min and

(SI1) 1% of range, linear decrease from 1 to 0; otherwise = 0 elevational suitability index if within species interquartile range for elevation, then = 1; else if between max/min and

(SI2) 10% of range, linear decrease from 1 to 0; otherwise = 0

continentality suitability if within species interquartile range for distance to ocean, then = 1; else if between max/

index (SI3) min and 20% of range, linear decrease from 1 to 0; otherwise = 0

biomic suitability index (SI4) = (percent presence of species in biome)/(highest percent presence of species in any biome) (ranges 0.01–1; values less than 0.01 are set to 0.01; 1 for biome with highest percent presence for the species)

biomic relative suitability = (percent presence of species in biome)/(percent presence of most common species in

index (SI5) biome) (ranges 0.01–1; values less than 0.01 are set to 0.01; 1 when most common species in biome)

1/5 habitat suitability index = (SI1 × SI2 × SI3 × SI4 × SI5) (ranges 0 to 1) (HSI) a Values for max/min range, interquartile range, and percent presence of species in a biome used in calculating SI1–SI5 are given in the inset tables of Figures 51–52. Values in bold are subjectively adjusted model parameters.

The values for five hand-fitted parameters of the HSI model (bold figures in above table) are used to assign marginal suitability to areas in which variable values (e.g., elevation) are outside of the range at which the species were found in the field in the Old World. Values for these five parameters are adjusted and various formulations for calculating HSI are evaluated (e.g., weighted arithmetic means; with or without SI5) in a

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 13 series of preliminary models of all five tamarisk beetles in order to minimize visually assessed errors regarding estimations of where several species are common in sympatry or where single species dominate in areas of either sympatry or nonsympatry in the final HSI models for both the Old World and (where introduced). The influence of various suitability indices and the inclusion of the biomic relative suitability index (SI5) on the sensitivity and elasticity of HSI models is analyzed using the following formulae to calculate sensitivity (S) and elasticity (E) (after Mitchell et al. 2002):

n n ∑()HSIi − HSI ∑()1− HSIi / HSI S = i=1 E = i=1 n , and n , HSI where is the mean HSI value from a series of n = 8 values of HSIi for which the value of a given suitability index ranges from 0.0 to 1.0 while all other suitability indices are held constant at 0.5. Suitability indices contributing to greater sensitivity and elasticity of HSI might be more important to include in the HSI model. A composite map displays where each Diorhabda spp. scores within the top 15% of the maximum HSI value of any Diorhabda species for a given grid cell (where HSI > 0.01).

Coden Institution AMA Azerbaijan Milli Akademiyasi, Baku, Azerbaijan BMNH Natural History Museum, London, United Kingdom CABP Commonwealth Agricultural Bureau Institute Bioscience Pakistan Centre, Rawalpindi, Pakistan DEI German Entomological Institute, Müncheberg, Germany HNHM Hungarian Natural History Museum, Budapest, Hungary MNMS National Natural Science Museum, Madrid, IRNSB Institut Royal des Sciences Naturelles de Belgique, Belgium, Brussels IZAS Institute of Zoology, Chinese Academy of Sciences, Beijing, China MRSN Regional Museum of Natural Science, Torino, Italy MSNM Civic Museum of Natural History, Milano, Italy MZHF Zoological Museum, University of Helsinki, Helsinki, Finland MZLU Museum of Zoology, Lund University, Sweden NHMB Natural History Museum, Basel, Switzerland NHRS Swedish Museum of Natural History, Stockholm, Sweden NMPC National Museum, Prague, Czech Republic NMSU New Mexico State University, Las Cruces, New Mexico, USA TAU Tel Aviv University, Tel Aviv, USNM National Museum of Natural History, Washington D.C., USA ZMAN Zoololgical Museum Amsterdam, Amsterdam, Netherlands ZMUH Universität von Hamburg, Zoologisches Institut und Zoologisches Museum, Hamburg, Germany ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany Individuals EGRC Edward G. Riley, College Station, Texas IAET Irfan A. Aslan, Erzurum, Turkey MBPF Michel Bergeal, Versailles, France RBCN Ron Beenen, Nieuwegein, Netherlands

14 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Label Data. All original label data from dissected and examined specimens are given under the listings of material examined for each species. Label data are ordered in a consistent manner; date formats are standardized and taxa on determination labels are abbreviated where appropriate. Additional information not on the labels, such as updated place names (from the GEOnet Names Server; NGA 2006) and geocoordinates are given in brackets. We also add in brackets the unique specimen label numbers placed on each dissected specimen and entered into a database of label data. An example of our specimen number label is “USDA USNM 2003-01”, where “USDA” is the name of our umbrella institution which is placed on each label (“USDA” is omitted from the label numbers in the lists), “USNM” represents an abbreviation designating the source collection, “2003” represents the year the specimen was dissected, and “01” represents a specimen number in our database. Data for examined voucher specimens from laboratory colonies are listed by their source field collection locations and includes receiver identifier codes (e.g., GSWRL-1999-8 [Temple, TX]) from invertebrate shipment records with foreign/overseas sources referenced in the USDA Agricultural Research Service (ARS) Germplasm Resources Information Network’s (GRIN) Releases of Beneficial Organisms (ROBO) database (USDA ARS 2006) under Invertebrate Germplasm information. Specimens. Most of the over 2,392 specimens examined (of which 784 were dissected) for this revision were loaned from the collections of the listed 21 institutions and four individuals. Vouchers of specimens from the USDA Grassland, Soil and Water Research Laboratory, Temple, Texas (GSWRL) are deposited at the USNM with the USDA-ARS Systematics Entomology Laboratory (SEL) under numbers GSWRL-2005-02 (for shipment and experimental vouchers) and GSWRL-2009-01 (North American field vouchers and general overseas collections).

Taxonomy

Genus Diorhabda Weise, 1883

Diorhabda Weise, 1883:316 (Type species: elongata Brullé, 1832, by original designation). Radymna Reitter, 1912:135 (Type species: Diorhabda rickmersi Weise, 1900, by monotypy). Prophyllis Reitter, 1912:135 (Type species not designated [undetermined hairy species]). Diorrhabda: Rusanov, 1949:118 (unjustified emendation).

Diagnosis. A future revision of the entire genus Diorhabda is needed, and it will probably involve additional transfers of current member species to other closely related genera of the tribe Galerucini Latreille (1802) (Galerucinae: Chrysomelidae), such as Galerupipla Maulik (1936) (see discussions below and under D. elongata group). After membership in the genus Diorhabda is stabilized, a description of distinguishing generic morphological characters can be made in comparison with other related genera. Included species. As of this revision, 14 species are recognized from the genus Diorhabda in Europe, Africa, Asia and Australia (New ) (Ogloblin 1936; Wilcox 1971, 1975; Berti and Rapilly 1973; Lopatin 1977a; Medvedev 1999). These 14 species are listed in the below discussion of information on their host plants under Biology. Discussion. Taxonomy. Galeruca elongata Brullé (1832) was made the type species for the genus Diorhabda by Weise (1883). Galerupipla persica (Faldermann, 1837) and G. fisheri (Faldermann, 1837), both of which attack Alhagi maurorum Medikus (Fabaceae) (Ogloblin 1936, Medvedev and Roginskaya 1988), were transferred from Diorhabda by Aslam (1972) without comment (see also Wilcox 1975), and this transfer has been widely overlooked (Lopatin 1977a, Mirzoeva 2001, Aslan et al. 2000, Lopatin et al. 2003, Warchalowski 2003, Gök and Duran 2004, Bieńkowski 2004, Lopatin et al. 2004). Other species transferred from Diorhabda include clarki Jacoby (1884; as D. robusta Jacoby [1899], Aslam 1972) and M. brevicornis (Jacoby, 1889) (Medvedev 1999). Both the genera Diorhabda and Galerupipla are in need of taxonomic revision. Our present revision is limited to treating the taxonomy of the five species of the Tamarix-feeding Diorhabda elongata group.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 15 TABLE 1. Host plant associations of the Diorhabda elongata group.a Species Host Plant Location Citation Diorhabda Italy Lundberg et al. 1987, present study elongata T. smyrnensis Greece (Crete), Turkey Regalin 1997; Gerling and Kugler 1973; Gök and Çilbiroğlu 2003, 2005; Gök and Duran 2004 T. hampeana Greece present study T. parviflora Greece, United States present study; R. Carruthers, pers. comm. (California) T. chinensis × T. United States (Texas) present study canariensis/T. gallica T. sp. Cyprus, , Greece Georghiou 1977, Tomov 1979, present study D. carinata T. ramosissima Georgia, Tajikistan, Lozovoi 1961, Kulinich 1962, present study Kazakhstan, Turkmenistan T. ramosissima/T. United States (Texas) present study chinensis T. smyrnensis (as T. Georgia, Azerbaijan Lozovoi 1961, Samedov and Mirzoeva 1985 hohenackeri) T. arceuthoides Tajikistan Kulinich 1962 T. hispida Tajikistan Kulinich 1962 T. aphylla Pakistan Habib and Hasan 1982, present study T. meyeri Azerbaijan Samedov and Mirzoeva 1985 T. c.f. indica (as T. cf. Pakistan present study troupii) T. aralensis Turkmenistan present study T. aucheriana Turkmenistan present study T. sp. Pakistan, Azerbaijan, Habib and Hasan 1982, Samedov and Uzbekistan Mirzoeva 1985, present study D. sublineata T. africana , Peyerimhoff 1926, Jolivet 1967 T. boveana (as T. Algeria Peyerimhoff 1926 bounopaea) T. gallica France, Spain Laboissière 1934, Hopkins and Carruth 1954 T. senegalensis present study T. aphylla Tunisia A. Kirk, pers. comm. T. spp. , Tunisia, Algeria, Boehm 1908; Alfieri 1976; S. Doguet, pers. Morocco comm.; present study Diorhabda T. ramosissima Uzbekistan, Kazakhstan, Sinadsky 1968; Mityaev and Jashenko 1998, carinulata China, United States 2007; DeLoach et al. 2003b; DeLoach et al. (Nevada, Utah, 2004 Wyoming, Colorado) T. hispida var. hispida Uzbekistan, China, Sinadsky 1968; Sha and Yibulayin 1993; Kazakhstan Mityaev and Jashenko 1998, 2007; DeLoach et al. 2003b continued.

16 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS TABLE 1. (continued) Species Host Plant Location Citation Diorhabda T. laxa China, Kazakhstan Sha and Yibulayin 1993; Mityaev and carinulata Jashenko 1998, 2007; DeLoach et al. 2003b (cont.) T. elongata China, Kazakhstan Sha and Yibulayin 1993; Mityaev and Jashenko 1998, 2007; DeLoach et al. 2003b T. arceuthoides China Sha and Yibulayin 1993, DeLoach et al. 2003b T. gracilis China, Kazakhstan Sha and Yibulayin 1993; Mityaev and Jashenko 1998, 2007 T. smyrnensis (as T. China Sha and Yibulayin 1993, DeLoach et al. 2003b hohenackeri) T. leptostachya Kazakhstan, China Mityaev and Jashenko 1998, 2007; present study T. kansuensis China Sha and Yibulayin 1993 T. androssowii China Sha and Yibulayin 1993 T. chinensis China, United States Sha and Yibulayin 1993, DeLoach et al. 2004 (Colorado) T. hispida var. karelinii China DeLoach et al. 2003b T. aralensis Turkmenistan present study poss. T. kotschyi (as T. Iran present study leptopetela)b T. parviflora United States (Nevada) Dudley et al. 2006; Dudley et al. in prep. T. sp. Kazakhstan, Mongolia, Mityaev 1958; Kulenova 1968; Lopatin 1970; China Medvedev and Voronova 1977a, 1977b, 1979; Medvedev 1982; Tian et al. 1988; Bao 1989; Ishkov 1996; Mityaev and Jashenko 1997, 1999, 2007; Li et al. 2000 Myricaria alopecuroides Mongolia Medvedev and Voronova 1979 M. sp. Mongolia, Kazakhstan Medvedev 1982; Mityaev and Jashenko 1997, 2007 D. meridionalis T. sp. Iran Berti and Rapilly 1973 poss. T. dioicab Iran present study aAll species referred to in previous citations as D. elongata, D. e. deserticola (China and Kazakhstan), or D. carinulata meridionalis (Iran). See text for each Diorhabda species under Distribution for discussion of data supporting corrected species identities. See Material Examined and Biology-Host Plants sections for details of new host records from the present study. Authors of plant names are given at their first mention in the text. b Possible hosts based on list of Tamarix and other plant species present at collection sites for the expedition (Hoberlandt 1981) with which Diorhabda we examined were collected.

Biology. Host Plants. The plant families attacked by different Diorhabda spp. range across the two subclasses Carophyllidae and Rosidae (plant taxonomic groupings according to Spichiger et al. 2004). Five species feed upon Tamarix (Carophyllidae: ): D. elongata, D. carinata, D. sublineata, D. carinulata and D. meridionalis (Table 1). Diorhabda carinulata additionally feeds upon Myricaria, also of the Tamaricaceae. Five other Diorhabda species also feed on plants of one or two closely related genera. Diorhabda rickmersi Weise (1900) feeds on Rumex sp. (Ogloblin 1936) and Rheum sp. (Medvedev and Roginskaya 1988) (both of Carophyllidae: Polygonaceae); D. lusca Maulik (1936) feeds on Celtis australis Linnaeus (Rosidae: Ulmaceae) (Maulik 1936); D. trirakha Maulik (1936) feeds on Ulmus wallichiana

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 17 Planchon (Ulmaceae) (Maulik 1936); D. rybakowi Weise (1890), which includes D. koltzei Weise (1900) as a junior synonym (Ogloblin 1936, Lopatin 1977a), feeds on Nitraria spp. (Carophyllidae: Nitrariaceae) (Medvedev and Voronova 1977b); and D. tarsalis Weise (1889) feeds on Glycyrrhiza uralensis Fischer et de Candolle (Rosidae: Fabaceae) (Medvedev and Voronova 1978). Hosts are unknown for four species of Diorhabda: D. inconspicua Jacoby (1894), D. nigrifrons Laboissière (1914), D. octocostata Gahan (1896), and D. yulensis Jacoby (1886).

Diorhabda elongata-Group tamarisk beetles

Diagnosis. The following diagnostic characters for tamarisk beetles are partly based upon keys to Diorhabda species for central Asia (Gressitt and Kimoto 1963a, Lopatin 1977a) and the Mediterranean region (Warchalowski 2003), and descriptions of certain south Asian Diorhabda species (Jacoby 1886, 1894; Maulik 1936). Tamarisk beetles may be distinguished from other species of Diorhabda by the following combination of characters: (1) the pronotum and elytra are glabrous or with very sparse setae, except for the pubescence of the epipleura; (2) the lateral borders of the prothorax are narrow, not flattened, with the posterior angles obtuse and distinct, not rounded or quadrate; (3) the elytral punctation is denser and finer than that of the pronotum; and (4) the elytron bears only a distinct lateral carina extending from the humeral calus. The description of external characters for adult D. carinulata (as D. e. deserticola) by Lewis et al. (2003b) applies generally to all species in the group, excepting measurements and coloration, which are discussed below in each species account along with genitalia. Included species. We recognize five sibling genitalic morphospecies of tamarisk beetles as forming the D. elongata group: D. elongata, D. carinata, D. sublineata, D. carinulata and D. meridionalis (Figs. 1–45). Distribution. The D. elongata species group is broadly distributed over the majority of the distribution of Tamarix from North Africa, southern Europe to central Asia (Map 1). Although tamarisk is also native to southern Africa, Israel and , and Southeast Asia (India, Bangladesh, and Burma) (Baum 1978), tamarisk beetles are unreported from these areas. Detailed distribution data with maps are discussed under the heading Distribution in each species account. Discussion. Taxonomy. Diorhabda elongata Brullé (as Galeruca) was originally described from the Pelopónnisos peninsula of Greece in 1832 (Brullé 1832). From 1858 to 1893, four other species described under Galeruca were synonymized under D. elongata (Reiche and Saulcy 1858, Weise 1893): (1) D. carinata (Faldermann, 1837) (of the Transcaucasus), (2) Galeruca sublineata (Lucas, 1849) (of Annaba, Algeria), which has been regarded as the subspecies D. e. sublineata (Lucas) (Gressitt and Kimoto 1963a), (3) D. carinulata carinulata (of Sarepta, Russia), and (4) G. costalis (Mulsant, 1852) (of southwest Turkey). We compare specimens of these four taxa collected from the vicinity of type localities to literature descriptions of type specimens. We similarly studied specimens corresponding to the subspecific taxa D. e. deserticola Chen (1961) (of Yuli, China; also examined paratypes), D. carinulata meridionalis Berti & Rapilly (of Minab, Iran), and D. e. carinata (Faldermann) (from central Asia) as regarded by Bechyné (1961). We characterize the variability of the genitalia and some external characters (size and elytral markings) for each taxon using specimens from the type localities (topotypes) where possible. We compare and match specimens of tamarisk beetles from throughout southern Europe, Africa and west and central Asia with the various taxa above, providing data on variability in genitalia across the entire distribution of each taxon. In our comparisons of the above taxa of tamarisk beetles, we examined male and female genitalia of 784 specimens from 37 countries. We distinguish five qualitatively distinct endophallic morphotypes based on endophallic sclerites. We also find geographically corresponding female genitalic morphotypes with distinct differences in internal sternite VIII and vaginal palpi. These five fully diagnosable genitalic morphotype pairs have additional nondiscrete differences in ranges of body length for each sex and, in some cases, striping of the elytra. Intermediate forms between these species are not seen in the extensive material examined, even in

18 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS several instances of sympatry and syntopy (sharing individual tamarisk trees as habitat). Most species appear to be at least parapatric or marginally sympatric and some are moderately sympatric. Among species with overlapping or abutting geographic ranges, the absence in nature of intermediate hybrid genitalic forms along a geographic cline towards areas of range contact constitutes a morphological hiatus that is inconsistent with a status of interbreeding subspecies (see Krysan et al. 1980, Patten and Unitt 2002) or geographic races. Although additional material would be desirable in the areas of abutting ranges of certain species such as in western Italy and and central Turkey, we believe the data available sufficiently document the lack of intermediate forms between species. The numbers of diagnostic genitalic differences between the few allopatric species are comparable with the number of diagnostic differences seen between moderately sympatric species. Consequently, a status of potentially interbreeding subspecies or races is also inconsistent for the few species living in allopatry (see Helbig et al. 2002). The maintenance of distinguishing genitalic characteristics in four species in culture under identical laboratory conditions is contraindicative of a status of intraspecific morphs, such as might occur in cases of genitalic polymorphism (e.g., Jocqué 2002) or phenotypic plasticity (e.g., Agrawal 2001). Distinct qualitative differences and morphometric discontinuities in genitalic structures distinguishing species are evidence of strong reproductive isolation in nature (Helbig et al. 2002). We find that these morphotype pairs are a complex of five fully diagnosable sibling genitalic morphological species (morphospecies) of Diorhabda. Further evidence for reproductive isolation between the four species D. elongata, D. carinata, D. sublineata, and D. carinulata is also found in differences in component ratios of putative aggregation pheromones (discussed below under Biology-Aggregation Pheromones) and reduced laboratory F2 hybrid egg viability (discussed below under Experimental Hybridization). We characterize the genitalia of D. elongata and corroborate the restoration of the species D. carinata and D. carinulata and their removal from synonymy with D. elongata by Berti and Rapilly (1973). Specimens identified by Bechyné as D. e. carinata are conspecfic with D. carinata. The following four taxonomic changes are made: (1) Galeruca sublineata Lucas is removed from synonymy with D. elongata and restored as the species D. sublineata (Lucas); (2) D. e. deserticola Chen is synonymized under D. carinulata; (3) D. koltzei ab. basicornis Laboissière is synonymized under D. carinulata;and (4) D. carinulata meridionalis Berti & Rapilly is elevated to species status as D. meridionalis Berti & Rapilly. The color variant D. e. ab. bipustulata Normand is synonymized under D. sublineata. Five sibling species (with no subspecies) are recognized as forming the D. elongata group: D. elongata, D. carinata, D. sublineata, D. carinulata and D. meridionalis. Each species of tamarisk beetle exhibits some degree of sympatry with at least one of the other species. Four of these five tamarisk beetle species were originally described in the 1800’s and one was described as a subspecies, but all have been identified and published under the name D. elongata by various workers (e.g., Wilcox 1971, Riley et al. 2003, Warchalowski 2003, Bieńkowski 2004, Lopatin et al. 2004). The five species of the D. elongata group may be distinguished by a combination of eight discrete, or near discrete, genitalic characters, involving the forms of the male endophallic sclerites and female vaginal palpi (Table 5), and the additional genitalic character of the form of female internal sternite VIII. Males of all species are distinguishable by the forms of one or more of three endophallic sclerites (PES, EES, and CES; Figs. 14–33). A combination of characters involving the forms of the vaginal palpi (VP) and internal sternite VIII (IS VIII) can be used to distinguish between females of three species, D. elongata, D. carinulata and D. meridionalis (Figs. 34–43). Females of two species, D. carinata and D. sublineata, are distinguished from the three other species by these same characters, but only in some cases are they distinguishable from one another, and only by the form of internal sternite VIII (Figs. 40–41). We find no differences in the morphology of the spermatheca (Figs. 34–38; SP) or median lobe (Figs. 14–18; ML) and tegmen (not shown) of the aedeagus that are sufficient for species diagnosis. Significant patterns in clinal geographic variation in the five species are not seen and, therefore, no subspecies are recognized. We provide keys for the sexing of adults, followed by keys to species for each sex based upon the genitalia.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 19 We are unable to distinguish the five sibling species of the D. elongata group solely on the basis of external characters. The presence of elytral vittae that extend into the basal half of the elytra (SMV and SSV; Figs. 5, 9) can be used to eliminate an identification of D. elongata, in which the elytral vittae, if present, are confined to the apical half of the elytra (Fig. 1). Differences in ranges of body length for each sex (Table 2) can aid in species identification of some individuals when used with distribution data (Map 1). Further investigation might reveal obscure external characters useful in separating some species such as external setation (e.g., Konstantinov and Lopatin 2000).

TABLE 2. External morphometrics ( ± SD [range], mm) of field collected material for the Diorhabda elongata species group.

Species n Length bodya Length left elytron Width elytra Ratio (LE)a (WE)b,c WE/LEa Males Diorhabda elongata 61 5.96 ± 0.38 b 4.63 ± 0.27 b 2.58 ± 0.19 c 0.56 ± 0.04 bc (5.33–6.78) (4.08–5.26) (1.56–3.04) (0.33–0.63) D. carinata 49 6.29 ± 0.50 a 5.01 ± 0.41 a 2.73 ± 0.21 b 0.55 ± 0.03 a (5.12–7.34) (4.01–5.81) (2.28–3.11) (0.48–0.60) D. sublineata 45 5.98 ± 0.50 b 4.67 ± 0.39 b 2.63 ± 0.31 c 0.56 ± 0.05 c (4.98–6.92) (3.74–5.47) (1.52–3.18) (0.29–0.62) D. carinulata 47 5.31 ± 0.35 c 4.16 ± 0.26 c 2.31 ± 0.21 d 0.55 ± 0.05 b (4.63–6.09) (3.74–4.84) (1.66–2.83) (0.36–0.61) D. meridionalis 14 5.12 ± 0.40 c 3.90 ± 0.32 c 2.18 ± 0.17 d 0.56 ± 0.03 bc (4.15–5.61) (3.11–4.29) (1.73–2.42) (0.52–0.63) Females D. elongata 32 6.60 ± 0.47 b 5.23 ± 0.38 b 2.96 ± 0.18 a 0.57 ± 0.02 bc (5.80–7.68) (4.71–6.09) (2.70–3.32) (0.52–0.60) D. carinata 31 6.76 ± 0.72 a 5.46 ± 0.59 a 2.97 ± 0.43 a 0.54 ± 0.06 a (5.05–8.44) (4.15–6.71) (1.63–3.74) (0.28–0.62) D. sublineata 24 6.52 ± 0.60 b 5.16 ± 0.46 b 2.94 ± 0.23 a 0.57 ± 0.03 c (4.91–7.40) (4.01–5.81) (2.49–3.39) (0.50–0.62) D. carinulata 23 5.80 ± 0.51 c 4.60 ± 0.49 c 2.53 ± 0.37 c 0.55 ± 0.04 b (4.98–6.99) (3.60–5.88) (1.28–3.25) (0.37–0.61) D. meridionalis 15 5.81 ± 0.44 c 4.51 ± 0.47 c 2.58 ± 0.19 c 0.58 ± 0.04 bc (4.77–6.37) (3.60–5.67) (2.15–2.84) (0.46–0.62) a Ranks of values followed by the same letter within the same column for each sex are not significantly different. The ranks of values for females are significantly larger than those of males with no sex by species interactions (P > 0.05; Two-Way Kruskal-Wallis Tests on ranks using PROC GLM-LSMEANS test; SAS Institute 2005). b Species by sex interaction is significant and ranks of values followed by the same letter within the same column (including both sexes) are not significantly different (P > 0.05; Two-Way Kruskal-Wallis Tests on ranks using PROC GLM-LSMEANS test; SAS Institute 2005). c Combined width of both elytra at widest point.

The endophalli (END) of the D. elongata group consist of an elongated tubular sac bearing one to three sclerites and various characteristic bulgings (Figs. 14–18). The sacs widen distally in D. elongata, D. carinata and D. sublineata, but narrow distally in D. carinulata and D. meridionalis (see Fig. 19d–e of Berti and Rapilly 1973 for an illustration of the fully exerted narrow tip of the endophallus in D. meridionalis). All species have a conspicuous palmate shaped dorsal sclerite, the palmate endophallic sclerite (PES), at the base of the sac that is armed subdistally or distally with one to several spines (or teeth) (Figs. 29–33). All species

20 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS also have an elongate shaped ventral sclerite, the elongate endophallic sclerite (EES), which runs along most of the length of the sac. The elongate sclerite also bears a raised blade which is armed with one to several spines (or teeth) (Figs. 19–23). The shapes of these two sclerites and the arrangement of the spines are useful in species diagnosis. Only D. sublineata posseses a third lateral sclerite that connects the palmate and elongate sclerites, the connecting endophallic sclerite (Figs. 16, 21, 31; CES). Our illustrations (Figs. 14–18) are of uninflated endophalli. Studies of inflated endophalli, such as obtained from mating pairs freshly killed by hot water, could reveal that the conformations of bulgings of the endophallus are also useful in species diagnosis. Scanning electron micrography is being done for the endophallic sclerites of the D. elongata species group by Jessica Perez, Roxana Reyna-Islas, and Dave Thompson at NMSU using some of our dissected material. Silfverberg (1974) found characters of the endophallic sclerites useful in helping differentiate between the genera Galerucella, , and in the tribe Galerucini. Endophalli in the D. elongata group closely resemble those of several Galerucella spp. (Figs. 1a, 4, 5 of Silfverberg 1974) which are also in the form of a tubular sac. Endophalli of Galerucella differ in bearing only a dorsal sclerite which is elongate, not palmate, in shape. The endophallus of D. lusca with its large serrated flagellum (Fig. 97 of Mann and Crowson 1996) is strikingly different from those in both the D. elongata group and Galerucella, and the placement of D. lusca in the genus Diorhabda should be reviewed. Spermathecae of the D. elongata group (Figs. 34–38; SP) more closely resemble those of some Galeruca spp. (Figs. 1d, 2d of Beenen 1999) than the more slender spermathecae of some Galerucella spp. (Fig. 6 of Hippa and Koponen 1986), but differ from both in bearing a pointed appendage at the distal tip (similar to that in North American , but more prominent; LeSage 1986). Additional studies are needed of the morphology of the endophalli and its sclerites and the female genitalia (vaginal palpi, spermathecae and internal sternite VIII; Figs. 34–38) in other species of Diorhabda and related genera. Such genitalic studies should be useful in any reviews of subtribal, generic, or specific classifications within the tribe Galerucini. Experimental Hybridization. D. C. Thompson et al. (in prep.) have conducted laboratory crossing studies between D. elongata and each of the three sibling species D. carinulata, D. sublineata, and D. carinata,and between D. carinulata and D. carinata (also Dave C. Thompson and Beth Peterson, New Mexico State University, Las Cruces, NM, pers. comm.). Additional crossing studies between D. elongata and D. carinulata have been conducted by some of us (DeLoach and Tracy unpublished data) and Julie Keller and Dan Bean (pers. comm.). These experiments are the subject of pending publications and involve interspecific pure line crossings, hybrid crossings, and concurrent pure line conspecific control crossings of laboratory reared virgin adult male and female beetles in no choice situations confined in vials with fresh tamarisk for food (n = ca. ten replications per type of cross). The mean percent viability of the F1 eggs produced from crossing the parental females and viability of the F2– F3 eggs from crossing progeny of the parental females were recorded. Diorhabda colonies used in these studies were found to be free of Wolbachia bacterial infections (D. Kazmer pers. comm.) that can reduce fertility in crosses between chrysomelid subspecies, such as is seen in the subspecies of Diabrotica virgifera LeConte (Giordano et al. 1997). In a no-choice laboratory environment, matings readily occurred between pure line D. elongata and the three other Diorhabda spp. (D. carinulata, D. carinata, and D. sublineata) and between D. carinulata and D. carinata, producing F1 hybrid eggs with 50–100% viability of control crosses, depending on parental species and their sex. In ♀ D. carinata × ♂ D. carinulata crossings, F1 hybrid egg viability was reduced by over 50% from that of pure line control crosses. But no F1 eggs were viable from ♀ D. carinulata × ♂ D. carinata crossings in which 90% of females died in copulo. A high incidence of female death in copulo also occurred in ♀ D. carinulata × ♂ D. elongata crossings and was associated with a 30% reduction in F1 egg viability over that of controls, but percent F1 egg viability was normal for ♀ D. elongata × ♂ D. carinulata crossings. Most surviving females failing to produce viable eggs in interspecific crosses probably failed to mate, but fertilization could also have produced inviable eggs. The larger size of male D. carinata and D. elongata, along with the larger more prominent distal basal facing spines of their palmate endophallic sclerites (Figs. 14–18), may be a factor in their observed difficulty disengaging from copulation with female D. carinulata and the associated high in copulo death rate for female D. carinulata.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 21 Reductions in F2 egg viability of ca. 50–100% from that of controls were found in at least one of the male/female crossing combinations for all interspecific hybrid/hybrid progeny crosses. Observed reductions in F1 and/or F2 hybrid egg viabilities is a form of hybrid breakdown (the opposite of hybrid vigor) providing further evidence for reproductive isolation between all the crossed Diorhabda species. Laboratory hybridization studies between the most morphologically similar species, D. sublineata and D. carinata, have not yet been conducted. However, these species strongly and asymmetrically differ with respect to the F1 and F2 hybrid egg viabilities resulting from the two combinations of male/female crosses of each species with D. elongata (D.C. Thompson and B. Peterson, pers. comm.), providing additional evidence of reproductive isolation. For example, in ♀ D. elongata × ♂ D. sublineata crosses, F1 egg viabilities are close to that of control crosses, but in ♀ D. elongata × ♂ D. carinata crosses, no F1 egg are viable. D. Thompson and B. Peterson (pers. comm.) found that hybrid breakdown in the form of reduced egg viability over that of controls is much less apparent in F2 and F3 progeny of laboratory produced hybrid/ hybrid crosses of D. sublineata × D. elongata (0–60% reductions) compared to that of F2 progeny produced from other hybrid/hybrid crosses in the D. elongata group (90% reductions). For example, egg viability of F2 D. elongata × D. carinulata hybrids is reduced by ca. 90–100% over that of pure line controls. However, egg viability of ♀ D. sublineata × ♂ D. elongata hybrids is reduced by about 60% from that of pure line controls for the F2 generation and it is reduced by about 50% for the F3 generation. In ♀ D. elongata × ♂ D. sublineata hybrids, egg viability is about the same as pure line controls in the F2 generation, but is reduced by about 25% in the F3 generation, possibly as a result of additional hybrid breakdown between generations. Colonies of D. sublineata × D. elongata hybrids can be easily maintained for at least six generations in the laboratory (D. C. Thompson and B. Peterson, pers. comm.) and F1 and F2 hybrids can have diagnostic anomalous hybrid character combinations (see D. sublineata × D. elongata Hybrid Morphology under species account for D. sublineata). Egg viability has never been measured in F4 or later generation hybrids, and, over several more generations of hybrid inbreeding, additional hybrid breakdown might lead to severe reductions in egg viability. Egg viability in backcross D. sublineata/D. elongata hybrids has not yet been characterized. With D. carinulata/D. elongata hybrids, egg viability appears normal when hybrid females are backcrossed with pure line males but there is no egg viability when hybrid males are backcrossed with pure line females (DeLoach and Tracy unpublished data). Additional forms of hybrid breakdown may occur in field situations (discussed below). Reduced F2 egg viability is also seen in laboratory hybrids of the chrysomelids carduorum Guerin- Meneville and A. cirsicola Ohno (Laroche et al. 1996). The chrysomelids Diabrotica longicornis (Say) and D. virgifera will also hybridize in the laboratory and produce fertile F1 and F2 hybrids. However, interspecific matings between these Diabrotica spp. are very rare in the field and prezygotic reproductive isolation apparently is operating through differences in adult feeding habits, pheromones, and phenologies (Hintz and George 1979). Interspecific laboratory crossings of sympatric populations of Galerucella nymphaeae and “G. sagittariae” from different host plants in Finland produce fertile F1 hybrids, but there is no cytological evidence of hybridization in nature (Nokkala and Nokkala 1998). The production of laboratory hybrids able to reproduce for several generations is not evidence for extensive hybridization and gene flow in nature (see Helbig et al. 2002). We have seen no morphological intermediates in nature that give evidence of hybridization between any species of the D. elongata group (for further discussion and characterization of male genitalia in laboratory produced hybrids, see D. sublineata × D. elongata Hybrid Morphology under D. sublineata species account). In addition, the genitalic morphology of laboratory produced male hybrids of D. sublineata and D. elongata cannot be attributable to other known species of Diorhabda. The lack of intermediate genitalic morphotypes between these species in nature, especially in areas of sympatry and parapatry, is strong evidence of reproductive isolation. Reproductive isolation in sympatric Diorhabda is probably maintained by prezygotic isolating mechanisms such as differing mate-recognition systems which may be reinforced by differences in aggregation pheromones (see Biology – Aggregation Pheromones below). It is possible that some of these Diorhabda species may hybridize rarely in nature and have limited flow between certain genes, as is seen in

22 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS some other closely related animal species (Helbig et al. 2002, Mallet 2005, Bull et al. 2006). For example, several species of passion-vine , Heliconius (: Nymphalidae: Heliconiinae), will interbreed and produce fertile hybrids in hybrid zones, but genetic analyses reveal strong barriers to interspecific gene flow, probably as a result of differing mate recognition systems (Jiggins et al. 1997, Bull et al. 2006). A limited amount of interspecific hybridization resulting in fertile hybrids also occurs between natural populations of some sympatric nonsibling Hawaiian species of Drosophila fruit (Diptera: Drosophilidae) which are strongly reproductively isolated by differing mate-recognition systems (Ahearn and Templeton 1989). The natural phenomenon of uncommon interspecific hybridization leading to fertile offspring is becoming more widely known among a variety of animal taxa (Mallet 2005). Diorhabda hybrids in the field might exhibit hybrid breakdown in areas additional to reduced egg viability, including behavioral disruptions in foraging (e.g., Godoy-Herrera et al. 1994) and pheromonal communication as a result of changes in hybrid pheromone composition (e.g., Wee and Tan 2005) and response (e.g., Lanier 1970). Laboratory or cage produced Diorhabda hybrids should not be considered as candidates for tamarisk biological control because of known and potential problems of hybrid breakdown in the field which would probably lead to low persistence and low efficacy. Further investigation is needed with genetic techniques to search for various types of hybrids and backcross hybrids and genetic introgression between Diorhabda species in the Palearctic. Morphometrics. Ranges for variables of all the external measurements overlap among males and females of all species in the D. elongata group (Table 2). Certain individuals of extremely small or large size can be identified as being of one or two species. For example, only females of D. carinata reach over 8.0 mm in length. Several significant differences in mean ranks of morphometric variables are seen. The mean lengths of the body and left elytron are significantly greater in females than males in each of the five species in the group. Mean lengths of the body and left elytron of both males and females of D. carinata are significantly greater than those of all other species in the D. elongata group (Table 2). Mean length of the body and left elytron are significantly smaller in males and females of D. carinulata and D. meridionalis compared to all other members of the D. elongata group. The combined width of the elytra at the widest point is significantly greater in females of D. elongata, D. sublineata, and D. carinata, than in females of D. carinulata and D. meridionalis and males of all species. The width of the elytra is significantly greater in male D. carinata compared to males of all other species and females of D. carinulata and D. meridionalis. The mean ratio of the width of the elytra to the length of the left elytron is significantly smallest in D. carinata making it the most elongate species in the group. Additional external morphometric variables are being evaluated for use in classification of the D. elongata group (Joaquin Sanabria, International Fertilizer Development Center [IFDC], Muscle Shoals, AL, pers. comm.). Milbrath and DeLoach (2006b) found the mean lengths of the left elytron (mm) in D. elongata (from Crete) reared on T. ramosissima × T. chinensis (4.57 ± 0.05 in 29 males, 5.19 ± 0.04 in 47 females) are slightly but significantly larger than that of individuals reared on T. aphylla (4.46 ± 0.04 in 42 males, 5.09 ± 0.05 in 33 females). These mean values for the length of the left elytron were similar to those observed for field material of D. elongata in this revision (Table 2), but the standard deviations they observed were much smaller than we observed. The wide geographic range of specimens measured in our revision probably accounts for the wider variability in measurements. Significant interspecific differences are seen in ranks of measurements of male endophallic sclerites (Table 3) and female vaginal palpi and internal sternite VIII (Table 4). All species of the D. elongata group significantly differ in the mean values of three variables: (1) the length of the blade of the elongate endophallic sclerite, (2) the ratio of the length of the blade of the elongate sclerite to the total length of the elongate sclerite, and (3) the width of the stalk of female internal sternite VIII. We find nine instances of discrete breaks or near breaks in size ranges of genitalic structures between species of the group, some of which are used in the keys to species (below): (1) both the range in width and width to length ratio of the palmate endophallic sclerite is nearly separate between D. carinulata and D. meridionalis, (2) the length of the elongate sclerite is largest in D. carinata and the range separates D. carinata from D. carinulata and D.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 23 meridionalis (see also scatter plot of Fig. 48); (3) the ratios of both the blade length and length of the spined area of the blade to the length of the elongate sclerite is smallest in D. elongata and the range separates D. elongata from all other species (see also scatter plot of Fig. 48), (4) the range in the ratio of the blade length to the length of the elongate sclerite separates D. carinulata and D. meridionalis (Fig. 48B); (5) both the range of width and width to length ratio of the vaginal palpus in D. carinulata and D. meridionalis is nearly separate from all other species in the group; (6) the range in width of the stalk of female internal sternite VIII separates D. meridionalis from all other members of the group except D. elongata; (7) the range in ratio of the width of stalk to width of apical lobe of female internal sternite VIII separates D. meridionalis from all other members of the group except D. elongata; and (8) the width of the lobe of the stalk in female internal sternite VIII separates D. meridionalis and D. carinata.

TABLE 3. Male genitalic morphometrics ( ± SD [range], mm) of field collected material for the Diorhabda elongata species group.a Palmate Endophallic Sclerite Elongate Endophallic Sclerite Species n Width (W) Length (L) Ratio W/L Length (L) Blade Length Ratio LB/L SpinedArea Ratio (LB) Length (SL) SL/L Diorhabda 27 0.31 ± 0.04 c 0.51 ± 0.08 c 0.61 ± 0.09 b 1.09 ± 0.06 c 0.37 ± 0.05 e 0.34 ± 0.05 e 0.05 ± 0.06 d 0.05 ± 0.05 c elongata (0.22–0.37) (0.38–0.73) (0.44–0.85) (0.96–1.23) (0.27–0.44) (0.24–0.42) (0.01–0.18) (0.01–0.16) D. carinata 21 0.40 ± 0.04 a 0.74 ± 0.07 a 0.54 ± 0.07 c 1.34 ± 0.08 a 0.67 ± 0.07 b 0.50 ± 0.05 d 0.58 ± 0.07 a 0.43 ± 0.05 b (0.32–0.46) (0.59–0.87) (0.40–0.69) (1.17–1.52) (0.57–0.82) (0.43–0.63) (0.46–0.68) (0.34–0.52) D. sublineata 22 0.33 ± 0.04 b 0.66 ± 0.06 b 0.50 ± 0.05 d 1.21 ± 0.10 b 0.62 ± 0.05 c 0.52 ± 0.03 c 0.52 ± 0.06 b 0.43 ± 0.05 b (0.27–0.40) (0.53–0.75) (0.42–0.63) (1.06–1.41) (0.52–0.72) (0.48–0.56) (0.38–0.64) (0.31–0.51) D. carinulata 23 0.34 ± 0.04 b 0.45 ± 0.04 d 0.77 ± 0.10 a 0.99 ± 0.07 d 0.56 ± 0.05 d 0.57 ± 0.05 b 0.44 ± 0.06 c 0.45 ± 0.05 b (0.28–0.43) (0.38–0.55) (0.61–1.02) (0.87–1.14) (0.46–0.63) (0.49–0.66) (0.34–0.54) (0.33–0.52) D. meridionalis 13 0.23 ± 0.02 d 0.49 ± 0.04 c 0.47 ± 0.07 d 1.03 ± 0.07 d 0.77 ± 0.06 a 0.75 ± 0.03 a 0.57 ± 0.08 a 0.55 ± 0.05 a (0.19–0.29) (0.41–0.56) (0.35–0.63) (0.87–1.12) (0.65–0.86) (0.67–0.79) (0.40–0.70) (0.46–0.64) a Ranks of values followed by the same letter within the same column are not significantly different (P > 0.05; Two-Way Kruskal-Wallis Tests on ranks using PROC GLM-LSMEANS test; SAS Institute 2005) (see Figs. 14–33 for illustrations of measured structures; see Fig. 48 for scatter plots of measurements of the elongate endophallic sclerite).

TABLE 4. Female genitalic morphometrics ( ± SD [range], mm) of field collected material for the Diorhabda elongata species group.

Vaginal palpi Internal sternite VIII Species n Length (LP) Width (WP) Ratio LP/WP Width stalk Width apical Ratio WST/ Width lobe of (WST) lobe (WAL) WAL stalk (WLS)b Diorhabda 29 0.16 ± 0.02 b 0.20 ± 0.01 b 0.78 ± 0.11 b 0.37 ± 0.06 d 0.70 ± 0.08 b 0.53 ± 0.10 c 0.08 ± 0.01 c elongata (0.11–0.19) (0.17–0.23) (0.52–0.94) (0.22–0.51) (0.52–0.84) (0.35–0.81) (0.06–0.11) D. carinata 21 0.16 ± 0.02 ab 0.22 ± 0.02 a 0.74 ± 0.12 bc 0.56 ± 0.07 b 0.82 ± 0.08 a 0.68 ± 0.10 b 0.14 ± 0.02 a (0.12–0.20) (0.19–0.26) (0.50–0.89) (0.42–0.68) (0.70–1.00) (0.53–0.86) (0.11–0.17) D. sublineata 19 0.17 ± 0.02 a 0.24 ± 0.03 a 0.71 ± 0.09 c 0.61 ± 0.07 a 0.79 ± 0.08 a 0.78 ± 0.06 a 0.12 ± 0.03 b (0.12–0.20) (0.18–0.30) (0.46–0.85) (0.52–0.77) (0.63–0.94) (0.62–0.87) (0.08–0.18) D. carinulata 20 0.17 ± 0.01 a 0.16 ± 0.01 c 1.06 ± 0.11 a 0.46 ± 0.07 c 0.71 ± 0.05 b 0.64 ± 0.09 b 0.14 ± 0.03 ab (0.14–0.19) (0.14–0.18) (0.94–1.36) (0.36–0.57) (0.60–0.81) (0.49–0.77) (0.07–0.22) D. meridionalis 12 0.18 ± 0.02 a 0.16 ± 0.02 c 1.13 ± 0.10 a 0.28 ± 0.03 e 0.68 ± 0.04 b 0.41 ± 0.05 d 0.06 ± 0.02 c (0.14–0.19) (0.13–0.17) (1.00–1.31) (0.22–0.33) (0.61–0.76) (0.33–0.48) (0.04–0.09) a Ranks of values followed by the same letter within the same column are not significantly different (P > 0.05; Two-Way Kruskal-Wallis Tests on ranks using PROC GLM-LSMEANS test; SAS Institute 2005). See Figs. 34–38 for illustrations of measured structures. b Width of widest lobe.

24 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS TABLE 5. Genitalic character state profiles for eight discrete, or near discrete, multi-state characters used in taxonomic keys of the Diorhabda elongata species group.a

Species Genitalic character Diorhabda elongata D. carinata D. sublineata D. carinulata D. meridionalis Elongate endophallic sclerite Increasing proportion of 0 0.5 0.5 0.5 1 blade length (length blade/ (0.24–0.42) (0.43–0.63) (0.48–0.56) (0.49–0.66) (0.67–0.79) length EES) Presence of hooked apex 0 0 0 0 1 (1); absence (0) Palmate endophallic sclerite Increasing spination of 0.5 1 1 0 0 distal margin (spination)b (1–6 [commonly 2–4] (2–6 [commonly (2–6 [commonly (1–2 subdistal (1–2 subdistal usually subdistal 4–5] usually distal 3–4] usually distal spines) spines) spines, maximum 2 spines, maximum 1 spines, maximum 1 spines distal) spine subdistal) spine subdistal) Increasingly narrowing 0.33 0 0 0.67 1 distal marginb (usually broadly (truncate serrate) (truncate serrate) (acutely (narrowly rounded, rarely rounded) rounded) truncate serrate) Presence of lateral 0 1 1 0 0 appendage (1); absence (0) Connecting endophallic sclerite Absent (0) vs. present (1) 0 0 1 0 0 Vaginal palpi Broadly rounded (0) vs. 0 1 1 0 0 narrowly rounded (1) Increasing elongation of 0 0 0 1 1 vaginal palpi (length palpi/ (0.52–0.94) (0.50–0.89) (0.46–0.85) (0.94–1.36) (1.00–1.31) width palpi)c aIncreasing degree or presence of a character state is denoted by a higher number. For details on diagnostic genitalic character states, see illustrated taxonomic keys, genitalic characters under each species, and Tables 2, 3 and 4. Character states are standardized to range from zero to one. Differences in character states between species are discrete with the exceptions noted below for three characters with near discrete differences for separating D. elongata from some species. bDifferences in spination and narrowing of the palmate endophallic sclerite are near discrete between D. elongata and the two species D. carinata and D. sublineata. cDifferences in elongation of vaginal palpi are near discrete between D. elongata and D. carinulata (difference is discrete if used in combination with width of widest lobe of female internal sternite VIII).

TABLE 6. Genitalic average taxonomic dissimilarity matrix (upper right) and genitalic Pearson product-moment correlation similarity matrix (lower left, shaded) for the Diorhabda elongata species group (from Table 5). a Species Species Diorhabda elongata D. carinata D. sublineata D. carinulata D. meridionalis D. elongata 0/1 0.571 0.672 0.449 0.680 D. carinata 0.200 0/1 0.354 0.746 0.884 D. sublineata 0.048 0.745 0/1 0.825 0.952 D. carinulata -0.007 -0.506 -0.703 0/1 0.412 D. meridionalis -0.115 -0.674 -0.944 0.728 0/1 a Matrices produced with NTSYSpc Interval Data (SIMINT) module (Rohlf 2006).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 25 Stenophenetic Analysis. The data matrix of genitalic character state profiles for the D. elongata group is shown in Table 5 and the derived average taxonomic dissimilarity matrix and a Pearson product-moment correlation similarity matrix are shown in Table 6. The genitalic dissimilarity and similarity phenograms produced from these matrices using the UPGMA clustering method (Figs. 49–50) were of the highest quality having the greatest rcoph values of 0.817 and 0.885, respectively. Genitalic phenograms produced from complete-linkage clustering and single-linkage clustering (not shown) have lower rcoph values of 0.745 and 0.811, respectively, for dissimilarity phenograms, and 0.882 and 0.862, respectively, for similarity phenograms. The genitalic similarity phenograms produced from any of the three clustering methods are of higher quality (have higher rcoph values) than genitalic dissimilarity phenograms produced from any of the clustering methods. Both UPGMA dissimilarity and similarity phenograms (Figs. 49–50) reveal D. carinata and D. sublineata are the most similar in genitalia, sharing seven of eight genitalic character states. D. carinata and D. sublineata form a group separate from the other three species in the genitalic dissimilarity phenogram while they form a group with D. elongata in the genitalic similarity phenogram. Diorhabda carinulata and D. meridionalis share five of eight genitalic character states and form a group with D. elongata in the genitalic dissimilarity phenogram but are separate from the other three species in the genitalic similarity phenogram. Similarities in genitalia among these Diorhabda species that are seen in the genitalic phenograms may or may not be correlated with genetic similarity. Berti and Rapilly (1973) originally described D. meridionalis as a subspecies of D. carinulata. Diorhabda sublineata differs from D. carinata by only a single genitalic character in each sex. Therefore, we suspect the D. carinulata/D. meridionalis and D. carinata/D. sublineata groupings do represent genetic similarity. The placement of D. elongata with either D. carinulata/D. meridionalis or D. carinata/D. sublineata has less support. A morphometric phenogram for the D. elongata group based upon external and genitalic data are in preparation (J. Sanabria, pers. comm.). Studies of genetic similarity in mitochondrial and nuclear DNA among D. elongata, D. sublineata, D. carinata, and D. carinulata are in progress (D. Kazmer, pers. comm.). Common Name. The vernacular name “tamarisk leaf beetle” has been applied in the Bulgarian, Russian and North American biological literature to D. elongata (Tomov 1974) as well as D. carinulata (as D. elongata: Sinadsky 1957, 1963, 1968; as D. e. deserticola: Bean et al. 2007b). The name “elongate tamarisk leaf beetle” was applied by Lozovoi (1961) to D. carinata (as D. elongata) in Georgia. The name “saltcedar leaf beetle” has been applied in North American biological control literature primarily to both D. elongata and D. carinulata (as D. elongata; DeLoach and Carruthers 2004b, Dudley and DeLoach 2004, Dudley 2005a, USDA Animal and Plant Health Inspection Service 2005, Carruthers et al. 2006, Dudley et al 2006, Herr et al. 2006, DeLoach et al. in prep., Moran et al. in press), but also to D. carinata and D. sublineata (as D. elongata; DeLoach and Carruthers 2004b). We propose the term “tamarisk beetles” to refer to all five species of the D. elongata group. Because Diorhabda are Old World species, in the common name we conserve the term “tamarisk”, used for the entire genus Tamarix in much of its native distribution (Baum 1978) and often in North America (e.g., Baum 1967a, Birken and Cooper 2006). In contrast, the competing term “saltcedar” is generally confined to use in the United States where it applies to certain weedy deciduous species (T. ramosissima/T. chinenis, T. canariensis/T. gallica, and T. parviflora; DiTomaso 1998). Members of the D. elongata group are probably the most important coleopterous defoliaters of tamarisk, making this group worthy of the general name “tamarisk beetle”. Consequently, the term “leaf” is dropped from the group vernacular name in order to shorten proposed common names for each species. Species common names are formed by adding a term referring to some unique intrinsic biogeographical or morphological characteristic before the term “tamarisk beetle”. Other species of the numerous beetles specializing on tamarisk outside the tribe Galerucini (Kovalev 1995) could be distinguished in their common name from “tamarisk beetle” by a name referring to their differing tribe (e.g., “tamarisk ” for Alticini), subfamily (e.g., “tamarisk casebearer” for Cryptocephalinae) or family (e.g., “tamarisk weevil” for Curculionidae). Vernacular names are listed following the names in the section header for each species account.

26 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Biology. Host Plants. Only and trees of the Old World family Tamaricaceae (order Tamaricales), including Tamarix and Myricaria, serve as hosts for any of the five species of tamarisk beetles (Table 1). Among tamarisks in North America, T. aphylla (Linnaeus) Karsten (athel tamarisk) is considered a non-target for biological control by Diorhabda because of its value as a shade tree and for windbreaks in parts of the southwestern U.S. and northern Mexico (DeLoach 1990; Map 7) and because it is only a minor invasive. However, T. aphylla is a major invasive along the Finke River in Australia (Griffin et al. 1989), and could potentially become more invasive in North America, where it is now known to invade by seed propagation in addition to vegetative propagation in southern Nevada (Walker et al. 2006) and northern Mexico (unpublished data; Map 7). Tamarix aphylla is a natural, but probably minor, host for D. carinata in Pakistan and D. sublineata in Tunisia (Table 1). In no-choice, small field cages, D. elongata, D. carinata, and D. sublineata, accepted T. aphylla for oviposition to the same degree as they accepted T. ramosissima × T. chinensis (Milbrath and DeLoach 2006b). In more sensitive multiple-choice, field cages studies, D. sublineata and D. carinulata generally prefer other North American tamarisks to T. aphylla for oviposition (egg-laying). However, D. elongata and D. carinata selected T. aphylla equally as well as other Tamarix species for oviposition in some tests, but oviposited on T. aphylla significantly less in other tests (Milbrath and DeLoach 2006a, 2006b), laying only ca. 22–30% as many eggs on T. aphylla as on some other tamarisks. In open field multiple-choice tests, D. elongata preferred ovipositing on T. chinensis × T. canariensis/T. gallica when compared with T. aphylla (Moran et al. in press). Tamarix aphylla appears to be at low to moderate risk to damage by the four tested Diorhabda spp. in the field. Potential damage by these Diorhabda to T. aphylla in a no-choice open field setting (especially where other Tamarix spp. are absent) is difficult to predict (Milbrath and DeLoach 2006b). The degree of potential damage to T. aphylla is still under investigation in field studies. Frankenia spp. (family Frankeniaceae) subshrubs and herbs are also in the order Tamaricales and occur throughout the native range of the D. elongata group (Jäger 1992) and in western North America (Whalen 1987). Native North American Frankenia spp. are of major concern as non-targets of tamarisk biological control whose safety must be insured (Lewis et al. 2003a; Milbrath and DeLoach 2006a; Herr et al. 2006, in prep.). Frankenia spp. can serve as a suitable but generally less favorable host in development from larvae to adult for D. elongata, D. carinata, D. sublineata and D. carinulata in caged no-choice studies (further discussed later; DeLoach et al. 2003b; Lewis et al. 2003a; Milbrath and DeLoach 2006a; Herr et al. 2006, in prep.). However, Frankenia spp. are never reported to serve as a hosts for any Diorhabda spp. under natural conditions in the open field (Table 1), even in specific surveys of Frankenia adjacent to where Diorhabda is found in Tunisia (DeLoach et al. 2003b). Three North American Frankenia spp., F. salina (Molina) I.M. Johnston, F. johnstonii Correll, and F. jamesii Torrey ex. A. Gray, provide almost no attraction for oviposition compared to Tamarix in field cage multiple-choice studies with D. carinulata (Lewis et al. 2003a, Milbrath and DeLoach 2006a) and D. elongata, D. carinata, and D. sublineata (Milbrath and DeLoach 2006a). In field cage no-choice studies with D. carinulata, D. sublineata and D. carinata, oviposition on F. jamesii and F. johnstonii was not different from non-host willow (Salix exigua Nutall) and adults experienced increased mortality compared to T. ramosissima × T. chinensis treatments (Milbrath and DeLoach 2006a). However, in laboratory cage no-choice tests, differences in oviposition by D. elongata on F. salina (inland variety) and T. ramosissima, T. parviflora, and T. aphylla were not significant (Herr et al. 2006). In open field testing of Frankenia salina transplanted among stands of Tamarix in Nevada and Wyoming, D. carinulata did not oviposit on F. salina which only sustained light (<4%) foliage loss from feeding of Diorhabda larvae that had crawled from nearby Tamarix (Dudley and Kazmer 2005). In open field tests with F. salina, T. ramosissima/T. chinensis and T. aphylla transplanted together in the field at Big Spring, Texas, D. elongata oviposited only a single egg mass on F. sa li n a compared with hundreds on T. chinensis × T. canariensis/T. gallica (Herr et al. 2006). Lewis et al. (2003a) and Milbrath and DeLoach (2006a) concluded that Frankenia is at very low risk of damage from any of the four tested Diorhabda species. Attack on non-preferred host plants by insect biological control agents is generally reduced as distance to the preferred host increases (Blossey et al. 2001). Consequently, we expect that damage by these four Diorhabda species to both Frankenia and T. aphylla is more unlikely the further away these plants are growing from preferred Tamarix

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 27 spp. hosts. Host preferences of D. meridionalis remain to be studied. Further information on the host range of the D. elongata group as well as additional biological data are reviewed later for each species. Phenology. Diorhabda carinulata has two to four generations from April through September and overwinter as adults in ground cover in western China and central Asia (Sinadsky 1968; Tian et al. 1988; Bao 1989; Sha and Yibulayin 1993; Mityaev and Jashenko 1998, 2007; Chen et al. 2000; Li et al. 2000. Diorhabda carinulata has two generations north of 38° in North America (DeLoach and Carruthers 2004b). Diorhabda elongata, D. carinata, and D. sublineata can have up to five generations from March to October in field cages at Temple, Texas (Milbrath et al. 2007). Five generations have also been observed for D. carinata (as D. elongata) at Ashgabat, Turkmenistan from March to September (Myartseva 1999). Diorhabda sublineata is collected from mid-January to mid-December in Tunisia (from examined material), where it may have more than five generations. The voltinism of D. meridionalis is yet to be studied, but adults are found from March to October in southern Iran (from examined material). Bean et al. (2007a) found the critical photoperiod (inducing diapause in 50% of the population) for adult D. carinulata from Fukang, China (44°N) is ca. 14 hours 55 minutes at 28°C. In comparison to D. carinulata from Fukang, the critical photoperiod for diapause induction is more influenced by temperature and much shorter in D. elongata from Crete (35°N), D. carinata from Qarshi, Uzbekistan (38°N), and D. sublineata from Sfax, Tunisia (35°N) (D. sublineata having the shortest critical photoperiod) (Bean and Keller in prep.). Bean and Keller (in prep.) also found intraspecific differences in critical photoperiod between two populations of D. carinulata from a similar latitude but widely varying elevations in China. Diorhabda carinulata from a low elevation at Turpan (42.8°N, -3 m elevation) have a critical photoperiod ca. 1 hr shorter than populations from Fukang (44.2°N, 552 m elevation) under laboratory conditions of ca. 25°C. These populations may represent two climatypes of D. carinulata adapted to differing seasonal progressions as influenced by elevation. Climatypes with intraspecific differences in critical daylength as influenced by seasonal differences across wide latitudinal and elevational gradients can probably be found among all the species of the D. elongata group. Additional observations on phenology are discussed in the species accounts. Aggregation Pheromones. Cossé et al. (2005) identified a male-produced aggregation pheromone in D. carinulata (as D. elongata) in laboratory and field studies. The pheromone consists of two components, the aldehyde (2E,4Z)-2,4-heptadienal and the alcohol (2E,4Z)-2,4-heptadien-1-ol. Robert Bartelt (USDA/ARS, Peoria, IL, pers. comm.) found that these same chemical components are also emitted by males of D. elongata, D. carinata and D. sublineata, but that the ratios of aldehyde to alcohol are not necessarily the same as in D. carinulata. In species other than D. carinulata, these chemicals are considered as putative pheromones until they can be field tested. The component ratio of alcohol to aldehyde in the putative pheromone of D. elongata of Sfakaki, Crete, Greece, is almost identical to that of D. carinulata. However, field testing revealed the D. carinulata pheromone is ineffective for D. elongata established at Big Spring, Texas (Allen Knutson, The Texas AgriLIFE Extension Service, Dallas, TX, pers. comm.). Diorhabda elongata from Posidi Beach, Greece are unusual in producing a higher proportion of alcohol in the pheromone, up to ca. 4 times higher than in the Crete population, as more pheromone is released. It is unclear whether the Crete D. elongata population may also be able to release higher amounts of pheromone under the proper conditions and prove to be more similar in pheromone composition to the Posidi Beach population than to D. carinulata (R. Bartelt, pers. comm.). If the Crete D. elongata pheromone is more similar to that of the Posidi Beach D. elongata, this would explain why the D. carinulata pheromone was ineffective for D. elongata. Proportions of alcohol in putative pheromones of D. carinata were intermediate between those of D. elongata from Crete and Posidi Beach. The proportion of alcohol in the putative pheromone of D. sublineata is about 10 to 20 times higher than that found in D. carinata, D. elongata from Crete, and D. carinulata, and about 5 times higher than that found in D. elongata from Posidi Beach. In some sympatric and syntopic species of bark beetles (Coleoptera: Scolytidae), pheromones of one species may disrupt conspecific pheromonal response of another species (Poland and Borden 1998), and the possibility of such competitive interactions between some sympatric and syntopic species of tamarisk beetles may warrant investigation.

28 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS TABLE 7. Biogeographic characteristics in the Diorhabda elongata species group.

Characteristic Diorhabda elongata D. carinata D. sublineata D. carinulata D. meridionalis Elevation (m) (location) Maximum 1,720 (Velez 2,903 (Sare Gearbid 2,606 (Tacchedirt, 1,834 (Bogda Shan 1,102 (Sekand, 27 Planina, Bosnia Mt., Afghanistan) Morocco) Mts., 80 km NW km ENE, Iran) Herzegovina) Urumqi, China) Minimum 1 (Lido di Volano, -15 (Sabirabad, 1 (LeBarcares, -55 (Turpan Botany 18 (Bilai, Iran) Italy) a Azerbaijan) France)a Station, China)

Latitude (°N) (location) Maximum 45.64 (Trieste 43.60 (Shelek 43.60 48.55 (Ynderbor, 36.20 (Halab, environs, Italy) environs, Kazakhstan) (Montpelier, Kazakhstan) Syria) France) Minimum 30.05 (Al Qahirah, 29.53 (Borazjan, 30 16.15 29.32 (Abareq, Iran) 25.73 (Bahu- Egypt) km NNE, Iran) (Ndiol Nar, Kalat, Iran) Senegal) Distance from ocean (km) (location) Maximum 255 (Malatya, 2,328 (Charyn River 503 (, Iraq) 2,282 (Burenhayrhan, 293 (Jolow Gir, Turkey) by Chundzha, Mongolia) Iran) Kazakhstan) Minimum 0 (Lido di Volano, 37 (Omidiyeh, 34 km 0 (LeBarcares, 266 (Abareq, Iran) 8 (Bilai, Iran) Italy) a SE, Iran) France)a

Major biomes inhabited (percentage of field collections)b Mediterranean Deserts & Xeric Mediterranean Deserts & Xeric Deserts & Xeric Forests, Woodlands (37.0%); Forests, Woodlands Shrublands (62.7%); Shrublands & Scrub (79.0%); Temperate Grasslands, & Scrub (73.2%); Temperate Grasslands, (62.5%); Temperate & Flooded Grasslands Savannas & Shrublands Temperate Broadleaf & Mixed Shrublands (31.5%); & Savannas (20.5%) Broadleaf & Forests (19.0%) Temperate Broadleaf (14.6%) Mixed Forests & Mixed Forests (25.0%) (16.3%) Biogeographic descriptorsc maritime temperate continental temperate maritime continental temperate maritime warm warm deserts/ subtropical cold and warm deserts/ subtropical Mediterranean grasslands/ broadleaf Mediterranean grasslands deserts/ broadleaf woodlands/ forests woodlands /flooded forests broadleaf forests grasslands aOne of several coastal locations with minimum values (See Figs. 51–52 for descriptive statistics of biogeographic characteristics summarized over 5 minute resolution grids). bBiomes ordered by percentage collections; biomes listed until cumulative percentage of collections from biomes equals at least 80% (Fig. 52B). cBased on descriptive statistics of biogeographic characteristics (Figs. 51–52).

Biogeography. General. The native range of the D. elongata species group is primarily restricted to the (Map 1). However, D. sublineata extends marginally into the in Senegal and and D. carinata extends marginally into the Indo-Malayan realm in northern Pakistan. All species of the D. elongata group are at least marginally sympatric with at least one other species in the group (Map 1; Table 8). Diorhabda carinulata and D. carinata are the most widely sympatric among the group, overlapping in distribution over a large portion of central Asia. The D. elongata group is distributed from 48°N in Kazakhstan (D. carinulata) to 16°S in Africa (D. sublineata) (Map 1, Table 7, Fig. 51B). Diorhabda carinulata is primarily found further north than all other species in the group while D. meridionalis is primarily found further south than all other species. The primary distributions of D. elongata, D. carinata, and

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 29 D. sublineata are intermediate in latitude within the D. elongata group, but D. sublineata is primarily found further south than D. elongata and D. carinata (Map1, Fig. 51B). All species may be found at or below sea level and D. carinata and D. sublineata can range to 2,903 and 2,606 m elevation, respectively (Table 7, Fig. 51A). Diorhabda elongata, D. sublineata,and D. meridionalis are primarily found at elevations below 400 m and in maritime regions. In contrast, D. carinata and D. carinulata are primarily found above 400 m and are more continental in distribution, occurring the furthest inland from the oceans (Map 1, Table 7, Figs. 51A, 52A).

TABLE 8. Biogeographic and habitat relationships among the Diorhabda elongata species group.a Species pairb Closest known points of range contact Closest Biogeographic/habitat distance relationship Diorhabda elongata/D. S. Turkey (Adana)–N. Syria (Halab) 186 kmc marginal sympatryc carinata D. elongata/D. Egypt (), S. Spain, Portugal, Algeria 0 km marginal sympatry sublineata* D. elongata/D. S. Russia (Dagestan Republic) 0 km?d marginal sympatry carinulata* D. elongata/D. S. Turkey (Adana)–N. Syria (Halab) 186 km parapatry meridionalis D. carinata*/D. Iraq (Baghdad) 0 km marginal sympatry sublineata D. carinata**/D. Azerbaijan, Turkmenistan, N. Iran, S. Uzbekistan, 0 km moderate sympatry, partial carinulata** Tajikistan, S. Kazakhstan, Kyrgyzstan, W. China syntopy

D. carinata**/D. W. Iran, Syria 0 km partial sympatry, partial meridionalis** syntopy D. sublineata/D. S. Azerbaijan (Ordubad)–C. Iraq (Baghdad) 635 km allopatry carinulata D. sublineata/D. W. Iran (Shush)–C. Iraq (Baghdad) 385 km allopatry meridionalis D. carinulata/D. S. Iran (Abareq–Sabzvaran) 102 km parapatry meridionalis a Closest point of range contact and distance are based upon examined material. In the text, we list locations where sympatric species were collected in close proximity, locations where syntopic species were collected from the same tamarisk trees or within the same series, and additional probable areas of range contact. See Map 1 for display of combined ranges of species. bSpecies with a single asterisk are dominant in known areas of weak or marginal sympatry with the other species in a pair. Species with two asterisks are sometimes dominant or sometimes less abundant than the other species in a pair, depending on the area (see text). c Actual closest distance between D. elongata and D. carinata may be zero based upon probable range contact between these species in E. Turkey (see text). dApproximate location (Dagestan Republic) of D. elongata specimen precludes measurement.

Eight desert, grassland and forest biomes characterize the biogeography of differing Diorhabda species (Table 9, Maps 2–6, Fig. 52B). Within these biomes, Diorhabda are most likely to be found in the primarily riparian, spring and maritime habitats of their host Tamarix species. The biomes inhabited vary widely among Diorhabda and host Tamarix species (Tables 9 and 11; Fig. 52B). Diorhabda elongata is usually collected in Mediterranean and Temperate Broadleaf forest biomes, and it is not reported from desert or grassland biomes as are many other species in the D. elongata group. The bioclimatic conditions to which D. elongata is adapted for Mediterranean maritime and riparian habitats are likely very different from conditions found in riparian habitats of the desert biome in which D. carinulata and D. meridionalis are primarily found.

30 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS TABLE 9. Biomic occurrence state profiles for degree of indigenous occurrence in eight biomes by country in the Diorhabda elongata species group.a

Biome Diorhabda D. carinata D. sublineata D. carinulata D. meridionalis elongata Deserts & Xeric Shrublands 0 1 1 1 1 (Afghanistan, (Algeria, Egypt, (Azerbaijan, China, (Iran, Azerbaijan, Iraq, Tunisia) Iran, Kazakhstan, Pakistan) Georgia, Iraq, Iran, Russia, Mongolia, Kazakhstan, Tajikistan, Pakistan, Turkmenistan, Tajikistan, Uzbekistan) Turkmenistan, Uzbekistan) Temperate Grasslands, 01 01 0 Savannas & Shrublands (Armenia, (Azerbaijan, Azerbaijan, Kazakhstan, Kazakhstan, Kyrgyzstan, Russia, Kyrgyzstan, Uzbekistan) Tajikistan, Turkey, Uzbekistan)

Montane Grasslands & 0 1 (Afghanistan, 0.5 1 0.5 Shrublands Pakistan, (Morocco) (China, Iran) (Iran) Tajikistan, Turkmenistan) Mediterranean Forests, 1 0.5 1 00.5 Woodlands & Scrub (Croatia, (Syria) (Algeria, France, (Syria) Cyprus, Greece, Morocco, Spain, Italy, Lebanon, Tunisia) Macedonia, Spain, Turkey) Temperate Broadleaf & 1 1 00.5 1 Mixed Forests (Bosnia (Azerbaijan, (Iran) (Iran) Herzegovina, Georgia, Iran, Bulgaria, Italy, Turkey) Macedonia, Turkey) Temperate Conifer Forests 0.5 1 00.5 0 (Turkey) (Iran, Turkey, (Iran) Pakistan) Flooded Grasslands & 0.5 01 00 Savannas (Egypt) (Algeria, Tunisia, Egypt) Tropical & Subtropical 0 0 0.5 00 Grasslands, Savannas & (Senegal) Shrublands Biomic Specialization Index 3.0 5.5 4.0 4.0 3.0 (BSI) aBiomes of Olson and Dinerstein (2002). States of occurrence in biome: 0—no record; 0.5—single record (minor presence); 1—multiple records (major presence).

Similarly, desert riparian habitats and grassland riparian habitats differ in bioclimatic and biogeographic characteristics. For instance, different salicaceous trees and shrubs generally characterize North American grassland riparian habitats, such as plains cottonwood (Populus deltoides Barton ex Marshall subsp. monilifera [Aiton] Eckenwalder), and black willow (Salix nigra Marshall), and various desert riparian habitats, such as Fremont cottonwood (P. fremontii Watson subsp. fremontii), Meseta cottonwood (P. fremontii Watson subsp. mesetae Eckenwalder), Rio Grande cottonwood (P. deltoides Barton ex Marshall subsp.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 31 wislizenii [Watson] Eckenwalder), and Goodding willow (S. gooddingii Ball) (Eckenwalder 1977, Powell 1998). Diorhabda carinata is also most often found in the desert biome but it occurs in the temperate grassland biome to a higher degree than other species of the D. elongata group. Like D. elongata, D. sublineata is also most common in the Mediterranean biome, but it also is common in the desert biome in which D. elongata is absent and has a strong presence in the Flooded Grasslands and Savannas biome in which D. elongata is uncommon (Fig. 52B). Diorhabda elongata and D. meridionalis occur in the fewest biomes and are the most stenobiomic species with low biomic specialization index (BSI) values of 3.0 (Table 9). Diorhabda carinata occurs in the largest number of biomes and is the most eurybiomic species, with the largest BSI value of 5.5.

TABLE 10. Biomic Bray-Curtis dissimilarity matrix for the Diorhabda elongata species group (from Table 9). a Species Species Diorhabda elongata D. carinata D. sublineata D. carinulata D. meridionalis D. elongata 0 ---- D. carinata 0.5040--- D. sublineata 0.568 0.555 0 - - D. carinulata 0.681 0.145 0.620 0 - D. meridionalis 0.500 0.276 0.409 0.409 0 a Matrix produced with NTSYSpc Interval Data (SIMINT) module (Rohlf 2006). Raw data matrix normalized with log (x + 1) transformation before dissimilarity matrix calculated.

Our knowledge of the biogeographic characteristics of the D. elongata group is limited by potentially under-collected areas such as eastern Turkey, Syria, Pakistan, Afghanistan and the . Additional comparative and descriptive information on the biogeography of each Diorhabda species, including the primary ecoregions they inhabit and their biogeographic associations with indigenous host Tamarix species, is discussed under the heading Biogeography in the species accounts. As an aid to matching Diorhabda species to target tamarisk species in North America, we compare the similarities in biomic profiles of the D. elongata group with those of tamarisk species invasive in North America below (see Biomic Analysis). Among these tamarisk species, T. aphylla and T. ramosissima are the most eurybiomic with BSI values of 8.0 and 6.0, respectively (Table 11). and T. parviflora are the most stenobiomic tamarisks with BSI values of 1 and 2.5, respectively. Tamarix austromongolica is closely related to T. chinensis (Zhang 2004), and our comparison of the native ranges and biomic profiles of these tamarisks prompted us to examine the potential presence of T. austromongolica in North America. Tamarix chinensis is widely cultivated throughout China, but it is indigenous only to eastern China (Zhang and Zhang 1990), east of 115°E in the Temperate Broadleaf and Mixed Forests biome. Tamarix chinensis has long been considered as widely invasive in the western U.S. (Baum 1967a; Gaskin and Schaal 2002, 2003), but the native biomic profile of T. chinensis does not include desert and grassland biomes from which it is reported in the western U.S. (Table 11). However, the native biomic profile of T. austromongolica does include desert and grassland biomes to which it is indigenous in the area between Lanzhou and Hohhot in north central China (Ma and Liu 1988; DeLoach, unpublished data; Zhang, Peng-yun, Lanzhou University, Lanzhou, China, pers. comm.; Map 5). Tamarix austromongolica was once considered a subspecies of T. chinensis (Zhou 1989), but recent studies support its status as a separate species (Hua et al. 2004, Zhang 2004). In response to our interest in the potential presence of T. austromongolica in North America, J. Gaskin haplotyped intron 4 of the nuclear phophoenolpyruvate carboxylase (pepC) gene (Gaskin and Schaal 2002) from specimens of cultivated T. austromongolica from China and found that some genetically match that of common T. chinensis and T. ramosissima/T. chinensis hybrids (J. Gaskin, pers. comm.). The fairly narrow distribution of T. austromongolica from ca. 103–112°E in

32 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS TABLE 11. Biomic occurrence state profiles for degree of indigenous occurrence in nine biomes by countries for Tamarix spp. invasive in North America.a

Biome T. T. austro- T. parviflora T. gallica T. T. aphyllac chinensis mongolicab canariensis Deserts & Xeric 1 01 00 1 1 Shrublands (Afghanistan, (China) (Algeria, (Afghanistan, Algeria, Azerbaijan, China, Morocco, Egypt, , India, Georgia, Iran, Iraq, Tunisia) Iran, Iraq, Israel, Kazakhstan, , Kuwait, Kyrgyzstan, Lebanon, , Mongolia, Pakistan, , Pakistan, Saudi Russia, , Arabia, Syria, Tunisia, Turkmenistan, Yemen) Uzbekistan) Temperate 1 01 0000 Grasslands, (Azerbaijan, China, (China) d Savannas & Iran, Iraq, Shrublands Kazakhstan, Kyrgyzstan, , Russia, Tajikistan, Turkmenistan, Ukraine, Uzbekistan) Montane 1 01 0001 Grasslands & (China, Kazakhstan, (China) (Afghanistan, Ethiopia, Shrublands Kyrgyzstan) Pakistan) Mediterranean 0001 1 1 1 Forests, (, (Algeria, (Algeria, (Egypt, Israel, Woodlands & Algeria, Croatia, France, France, Italy, Lebanon, Libya, Scrub France, Greece, Italy, Libya, Morocco, Palestine, Italy, Libya, Morocco, Morocco, Tunisia) Portugal, Spain, Spain) Portugal, Turkey) Spain, Tunisia) Temperate 1 1 01 1 1 1 Broadleaf & (Armenia, (China) (Greece, Italy, (France, (Spain) (Iran) Mixed Forests Azerbaijan, China, Portugal) Italy, Spain, Georgia, Iran, Iraq, United Russia, Turkey) Kingdom) Temperate 1 0 0 0.5 0.5 01 Conifer Forests (China, Kazakhstan, (Turkey) (Morocco) (Afghanistan, Iran, Kyrgyzstan, Pakistan) Pakistan) Flooded 1 0 0 0 0 0.5 1 Grasslands & (Iraq) (Libya) (Egypt, Iraq, Tunisia) Savannas Tropical & 00000.501 Subtropical (Senegal) (Ethiopia, , Grasslands, ) Savannas & Shrublands Tropical and 0000001 Subtropical Dry (India) Broadleaf Forests Biomic 6 1 3 2.5 3 3.5 8 Specialization Index (BSI) aBiomes of Olson and Dinerstein (2002). States of occurrence in biome: 0—no record; 0.5—single record (minor presence); 1—multiple records (major presence). b Invasiveness in North America requires further confirmation. c Currently minor invasive. d Distribution based on hand-drawn map of Zhang Peng-yun (pers. comm.).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 33 China is situated at the eastern edge of the distribution of T. ramosissima and approaching the westernmost edge of the distribution of T. chinensis (115°E) where a T. ramosissima/T. chinensis hybrid zone would be expected to occur (Map 5). In view of the biomic profile, DNA data, and distribution of T. austromongolica, the possibilities that this species may be a T. ramosissima/T. chinensis hybrid and that it is contributing to the genome of invasive T. ramosissima/T. chinensis in western North America warrant further investigation. Biomic Analysis. The biomic profile for the D. elongata species group (Table 9) is used to generate a biomic Bray-Curtis dissimilarity matrix (Table 10) in order to compare similarities in biomic profiles within the group. Of the several clustering methods used to generate biomic dissimilarity dendrograms from the dissimilarity matrix, the UPGMA clustering method produces the biomic dendrogram with the greatest rcoph value of 0.879 (Fig. 53). Biomic dendrograms produced from complete-linkage clustering and single-linkage clustering (not shown) have lower rcoph values of 0.850 and 0.855, respectively. In the UPGMA biomic dendrogram, D. carinata and D. carinulata are the most similar in biomic profiles. The close relationship in biomes inhabited by D. carinata and D. carinulata can be related to these species having the greatest degree of sympatry and syntopy in the D. elongata group (Table 8, Map 1). Diorhabda meridionalis is partially sympatric with D. carinata and is grouped next followed by D. sublineata. Diorhabda elongata is grouped separately from all other tamarisk beetles. We also test for a positive correlation in the dissimilarities in biomic profiles of the D. elongata group (Table 10) to dissimilarities in key morphological characters of the genitalia (Table 6, discussed previously). The biomic Bray-Curtis dissimilarity matrix of the D. elongata group (Table 10) is not significantly positively ≥ correlated with the genitalic average taxonomic dissimilarity matrix (Table 6) (rM = -0.40559, P[random rM observed rM] = 0.9211 with 9,999 permutations, Mantel Test; NTSYS Matrix Comparison Plot [MXCOMP] module [Rohlf 2006]). In other words, we find no significant positive correlation between the similarity in biomes inhabited by the five Diorhabda species and similarities in their genitalia. Instead, genitalic similarity tends to be negatively correlated with biomic similarity. This negative correlation is seen in species pairs such as D. carinata/D. carinulata, which are similar in biomic profiles but dissimilar in genitalic character profiles, and D. carinata/D. sublineata, which are different in biomic profiles, but similar in genitalic character profiles (Figs. 49 and 52; see Discussion—Stenophenetic Analysis above). An exception to genitalically similar species more greatly differing biomically is found in the species pair D. carinulata/D. meridionalis, which are both common in the Deserts and Xeric Shrublands biome. However, D. carinulata and D. meridionalis are parapatric (Map 4), and the apparent biogeographic distinctiveness of these species probably results from different preferences for latitude and distance to the ocean within the desert biome (discussed later; see Figs. 51–52). Further ecogeographic studies in the D. elongata group should include testing for a negative correlation between morphological similarity and a profile of a variety of climatic variables. The combined biomic profiles of the D. elongata group (Table 9) and tamarisks invasive in North America (Table 11) are used to generate a Bray-Curtis dissimilarity matrix (Table 12) in order to assess similarities in biomic profiles among Diorhabda and Tamarix together. Two biomic dissimilarity dendrograms from the dissimilarity matrix that are produced using the UPGMA clustering method have the greatest rcoph value of 0.881 (one dendrogram is shown in Fig. 54). Several biomic dendrograms produced from complete- linkage clustering and single-linkage clustering (not shown) have lower rcoph values of 0.841 and 0.808, respectively. The two UPGMA biomic dendrograms differ only in alternating the places of T. gallica and D. elongata (Fig. 54). In these dendrograms, D. carinulata, D. carinata, T. ramosissima, and T. austromongolica form a group similar in biomic profiles. Diorhabda elongata, T. parviflora, and T. gallica also form a group associated with T. chinensis. Diorhabda meridionalis and T. canariensis form a group from which D. sublineata occurs next up the tree. In order to display spatial relationships in biomic profiles among the Diorhabda and Tamarix species, a three dimensional biomic principle coordinate analysis (PCoA) scatter plot (Fig. 55) is computed from the biomic Bray-Curtis dissimilarity matrix for Diorhabda and Tamarix (Table 12). Visual groupings of the Diorhabda and Tamarix species generally coincide with relationships in the biomic dissimilarity dendrogram

34 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS (Fig. 54). The two groupings of D. elongata/T. gallica/T. parviflora and D. meridionalis/T. canariensis are the most distinct in the biomic PCoA scatter plot. The group of D. carinata/T. ramosissima/D. carinulata/T. austromongolica is also evident, but less distinct. The taxa of D. sublineata and T. aphylla are more isolated as in the biomic dendrogram. The influence or loadings of the different biomes on PCoA axes are computed as Spearman rank-order coefficients (Table 13—Spearman rank-order correlation coefficients) for ranks of species in biomes (from Tables 9 and 11) versus ranks of species in PCoA axes (from Table 13—Eigenvectors). The D. elongata/T. gallica/T. parviflora group has high eigenvectors along PCoA axis one (Table 13—Eigenvectors, Fig. 55) which is strongly negatively correlated with the Desert and Xeric Shrublands, Temperate Grasslands, Savannas and Shrublands, and Montane Grasslands and Shrublands biomes (Table 13- Spearman rank-order correlation coefficients), but positively correlated for Mediterranean and Temperate Broadleaf and Mixed Forests biomes. In other words, deserts and grasslands are strongly contraindicative for the presence of species favoring Mediterranean and Temperate Broadleaf and Mixed Forests biomes, such as D. elongata, T. gallica and T. parviflora. In contrast, eigenvectors for D. carinulata, D. carinata, T. ramosissima, and T. austromongolica, are very low along axis one as these species have a strong presence in deserts and temperate and montane grasslands. Eigenvectors for T. aphylla and D. sublineata are highest along PCoA axis two which is strongly positively correlated with Flooded Grasslands and Savannas. The eigenvector for D. carinata is the highest along axis three which is strongly positively correlated with the Temperate Conifer Forests biome.

TABLE 12. Biomic Bray-Curtis dissimilarity matrix for the Diorhabda elongata species group and invasive North American Tamarix species (From Tables 9 and 11).a

Species Species D. elongata D. carinata D. sublineata D. carinulata meridionalis D. T. ramosissima chinensis T. T. austromongolica T. parviflora T. gallica T. canariensis T. aphylla

Diorhabda elongata 0.000 D. carinata 0.188 0.000 D. sublineata 0.184 0.217 0.000 D. carinulata 0.213 0.116 0.189 0.000 D. meridionalis 0.149 0.154 0.154 0.143 0.000 Tamarix ramosissima 0.209 0.142 0.197 0.116 0.188 0.000 T. chinensis 0.130 0.224 0.201 0.188 0.130 0.224 0.000 T. austromongolica 0.239 0.174 0.170 0.083 0159 0.174 0.201 0.000 T. parviflora 0.059 0.179 0.206 0.205 0.137 0.288 0.116 0.232 0.000 T. gallica 0.083 0.188 0.197 0.213 0.149 0.236 0.130 0.239 0.059 0.000 T. canariensis 0.116 0.183 0.143 0.197 0.093 0.205 0.154 0.209 0.130 0.143 0.000 T. aphylla 0.209 0.201 0.188 0.232 0.209 0.201 0.265 0.265 0.228 0.209 0.205 0.000 a Matrix produced with NTSYSpc Interval Data (SIMINT) module (Rohlf 2006). Raw data matrix normalized with log (x + 1) transformation before dissimilarity matrix calculated.

Habitat Suitability Index Models. Descriptive statistics of the four biogeographic variables (biome, latitude, elevation and distance to ocean) for each Diorhabda species are displayed in Figures 51–52. These four descriptive statistics are used in calculating the five suitability indices (SI1–SI5; Figs. 56–58), and the final habitat suitability index (HSI; the geometric mean of SI1–SI5) in hand-fitted HSI models for each Diorhabda species We subjectively adjusted parameters of the suitability indices in preliminary HSI models

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 35 (not shown) to reduce the visually assessed overall error in the final models (see Materials and Methods). Our sensitivity and elasticity analysis of the five suitability indices revealed that the categorical biomic indices SI4 alone and both SI4 and SI5 together produce significantly higher model elasticities than do the continuous linear variable indices SI1, SI2, and SI3 (Table 14).

TABLE 13. Data from principal coordinate analysis (PCoA) of biomic Bray-Curtis dissimilarity matrix (Table 12) for species of Diorhabda and Tamarix. a

Principal Coordinate Eigenvector Axis Category 1 2 3 4 5 6 Species Eigenvectors Diorhabda elongata 0.093 0.010 0.020 0.025 0.020 0.031 D. carinata -0.062 0.015 0.086 -0.053 0.003 0.019 D. sublineata -0.021 0.032 -0.116 0.007 -0.006 0.037 D. carinulata -0.102 -0.047 0.026 0.003 -0.015 -0.009 D. meridionalis 0.008 -0.029 -0.023 -0.046 0.017 -0.046 T. ramosissima -0.105 0.027 0.040 0.063 0.049 0.002 T. chinensis 0.069 -0.083 -0.011 0.048 -0.009 -0.040 T. austromongolica -0.111 -0.078 -0.038 -0.010 -0.034 0.015 T. parviflora 0.098 -0.020 0.039 -0.005 -0.010 0.015 T. gallica 0.100 0.004 0.027 -0.001 -0.042 0.012 T. canariensis 0.050 0.007 -0.042 -0.038 0.061 -0.003 T. aphylla -0.017 0.163 -0.008 0.006 -0.034 -0.033 Eigenvalue 0.074 0.045 0.030 0.014 0.011 0.008 % Axis Loading 39.992 24.415 16.155 7.355 6.111 4.438 % Cumulative Axis Loading 39.992 64.407 80.561 87.916 94.028 98.881 Spearman rank-order correlation coefficients for ranks of species in biomes (from Biome Tables 9 and 11) versus ranks of species in PCoA axes (from above eigenvectors)

Deserts & Xeric Shrublands -0.819* 0.307 -0.256 -0.307 0.205 -0.154 Temperate Grasslands, Savannas & -0.819* -0.102 0.410 -0.102 -0.051 0.102 Shrublands Montane Grasslands & Shrublands -0.899* 0.238 0.185 -0.079 -0.159 -0.079 Mediterranean Forests, Woodlands & 0.618* 0.473 0.191 -0.366 0.069 0.321 Scrub Temperate Broadleaf & Mixed Forests 0.580* 0.161 0.470 -0.023 0.354 -0.304 Temperate Conifer Forests -0.112 0.551 0.834* 0.156 -0.052 0.138 Tropical and Subtropical Dry Broadleaf -0.044 0.480 -0.044 0.131 -0.306 -0.306 Forests Flooded Grasslands & Savannas -0.181 0.816* -0.225 0.524 0.402 0.189 Tropical & Subtropical Grasslands, 0.170 0.557 -0.184 0.198 -0.479 0.078 Savannas & Shrublands

a PCoA eigenvectors and eigenvalues computed (NTSYSpc Eigenvetors [EIGEN] module; Rohlf 2006) from double centered symmetric dissimilarity matrix (not shown; NTSYSpc Dcenter module). Spearman rank-order correlation coefficients with asterisk are significant (P < 0.05; Proc Corr; SAS Institute 2005). See Figure 55 for biomic PCoA scatter plot of the first three axes.

36 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Our final HSI models for the D. elongata group in the Palearctic (Maps 8 and 9) are generally accurate in estimating the optimal native range for each Diorhabda species. However, the models overestimate the optimal native ranges in some cases such as for D. elongata in France, Spain, and northwest Africa where D. sublineata dominates (Map 8a) and for D. sublineata in southeastern Europe from which only D. elongata is known (Map 8c). The domination of either D. elongata or D. sublineata in parapatric areas of similar suitability could be related to potentially strong competitive interactions between these species.

TABLE 14. Sensitivity and elasticity ( ± SD) of habitat suitability indices (HSI) for the Diorhabda elongata species group as influenced by suitability indices (SI) for HSI models with and without the biomic relative suitability index a (SI5). Sensitivity Elasticity

Suitability index n HSI models without SI5 HSI models with SI5 HSI models without SI5 HSI models with SI5

SI1, SI2 or SI3 (D. 8 0.131 ± 0.132 a 0.122 ± 0.134 a 0.303 ± 0.306 a 0.281 ± 0.310 a elongata group)

SI4 or SI4/SI5 Diorhabda 8 0.118 ± 0.087 a 0.163 ± 0.118 a 0.435 ± 0.322 b 0.686 ± 0.495 b elongata D. carinata 8 0.127 ± 0.082 a 0.184 ± 0.090 a 0.328 ± 0.211 b 0.490 ± 0.238 b D. sublineata 8 0.118 ± 0.076 a 0.154 ± 0.113 a 0.404 ± 0.258 b 0.556 ± 0.409 b D. carinulata 8 0.133 ± 0.065 a 0.190 ± 0.084 a 0.433 ± 0.210 b 0.652 ± 0.289 b D. meridionalis 8 0.123 ± 0.082 a 0.181 ± 0.109 a 0.395 ± 0.264 b 0.650 ± 0.391 b Totalb 160 0.129 ± 0.114 a 0.135 ± 0.122 a 0.311 ± 0.279 a 0.363 ± 0.340 a a Means of a given variable and the same sample size followed by the same later are not significantly different (P > 0.05; Fisher’s Protected LSD Test using PROC GLM-LSMEANS test; SAS Institute 2005). Sensitivities and elasticities are calculated across eight possible values ranging from 0.0 to 1.0 for the analyzed suitability index while the other indices are held constant at 0.5. For the continuous variables SI1, SI2 and SI3, eight values are chosen at equal intervals of 0.125

(0.0, 0.125, 0.250,…1.0) and these do not vary between species. For the categorical variables SI4 and SI5, the eight fixed possible values (from Figs. 57B and 58) vary between species, and SI4 and SI5 vary directly with one another and are considered together in calculating sensitivity and elasticity. b Total sample size (n) = five species × four indices × eight values per index = 160.

The estimated North American ranges of the various Diorhabda spp. vary widely (Maps 10 and 11). The HSI model for D. elongata correctly predicts both the high suitability of the Cache Creek, California site and the low suitability of the Artesia, New Mexico and Lake Meredith, Texas sites. However, our D. elongata HSI model underestimates the suitable habitat at sites in west Texas at Big Spring, Imperial, and Pecos, where D. elongata has established well but where the model continentality suitability index and biomic suitability indices are near zero (Map 10a). The HSI model for D. carinulata correctly predicts the high suitability of northern desert habitats in Nevada, Colorado and Wyoming, and additionally predicts suitable habitat in the northern . Our HSI models estimate that D. carinata should have the widest area of suitable habitat in North America (Map 10b), while D. elongata is estimated to have the smallest area (Map 10a). The wider area of suitability for D. carinata is related to its being the most eurybiomic species (see Biogeography—General above), giving it a wide tolerance in the biomic suitability indices (Figs. 57B and 58), and having a relatively wide tolerance in both the elevational and continentality suitability indices (Figs. 56B and 57A). The smaller estimated area of suitability for D. elongata in North America is related to it being among the most stenobiomic species, giving it a smaller tolerance for the biomic suitability indices (Fig. 57B and 58), and having a very narrow tolerance in the continentality index. D. sublineata is estimated to have a similar area of optimal suitability to D. elongata in Mediterranean biome along the Pacific coast, but has a

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 37 higher suitability for southern desert biomes (Map 10c). D. meridionalis is predicted to have the highest suitability in southern maritime desert habitats such as the Sonoran Desert and Tamaulipan Mezquital in southern Texas (Map 11b). The composite maps displaying which Diorhabda spp. score within the top 15% of the maximum HSI value of any Diorhabda spp. are generally accurate in estimating which single species or multiple species dominate in any given area of their native range (Map 12). For example, both the dominance of D. carinulata in China and the northern part of its range, and the dominance of D. carinata is in some grasslands of Central Asia are depicted fairly accurately. The composite HSI maps fail to depict the dominance of D. sublineata in France, Spain, and the coast of northwest Africa and the dominance of D. elongata in the northeastern Mediterranean, instead depicting both species as equally dominant in these areas. The composite HSI maps for the D. elongata group for North America (Map 13) estimate D. carinulata as dominant or co-dominant with D. carinata in areas where it has established well above 38°N in temperate cold deserts such as the Great Basin Steppe, Colorado Plateau Shrublands and Wyoming Basin Shrub Steppe. Diorhabda carinulata and D. carinata are estimated as potentially dominating or co-dominating in temperate warm desert areas areas below 38°N such as the northern Mojave Desert, southern Colorado Plateau Shrublands, and the Trans-Pecos region of the Chihuahuan Desert. Diorhabda carinata is estimated to dominate in large areas of temperate grasslands in the Great Plains, such as the Western Short Grasslands (including where D. carinulata has established at Pueblo, Colorado) and Central and Southern Mixed Grasslands, and in temperate conifer forests, such as the Arizona Mountains Forests in Arizona and New Mexico (Map 13). Although D. carinulata is established in the Western Short Grasslands at Pueblo, Colorado, our model predicts that D. carinata is better suited to this habitat for which it is in a more optimal range in terms of biome (being a grassland), latitude, and distance to the ocean. Pueblo, Colorado falls within the HSI environmental envelope suitability score (SI5) for D. carinulata, but the model map only displays D. carinata for Pueblo because this species is more than 15% higher in the overall optimality (SI10) score. Introduction of southern climatypes of D. carinulata and D. carinata may speed their adaptation to areas south of 34°N (see further discussion in species accounts under Potential in Tamarisk Biological Control). Diorhabda elongata is correctly estimated as dominating the Mediterranean biome of northern California, including where it is established vigorously at Cache Creek. However, our models estimate that areas of west Texas, where D. elongata has also established well, are more suitable for D. carinata and D. carinulata. West Texas falls outside the environmental envelope suitability scoring (SI5) for D. elongata in both biome (being in temperate grasslands) and distance to ocean limits (over 600 km from ocean versus a 255 km limit) (Table 7, Fig. 52). The environmental envelope suitability score portion of our HSI model underestimates the potential range of D. elongata in grasslands and desert of west Texas, but our overall HSI model score may still be accurate in predicting that D. carinata and D. carinulata are better suited to these areas. Diorhabda sublineata is estimated as dominating in the Mediterranean biome in California below 37°N and the Chihuahuan Desert below 29°N. Diorhabda meridionalis, a species with a higher preference for maritime subtropical deserts between 26–31°N, is estimated as dominating over most of the southern Sonoran desert and Tamaulipan Mezquital xeric . We consider the estimations of these hand-fitted HSI models as first rough approximations of the optimal tamarisk beetle species for various Old World and New World tamarisk habitats. The descriptive statistics used in calculating HSI grids could change substantially for some Diorhabda species with additional distribution data from under-collected areas. Complex interactions of the biogeographic variables we examined between both one another and unexamined bioclimatic variables probably occur, especially near environmental extremes for each species or in novel environments such as North America. Bioclimatic variables are among the best predictors of species distributions and further studies of the distributions of tamarisk beetles are planned using climatic data with species distribution models designed for presence-only data (e.g., Elith et al. 2006).

38 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS More detailed comparisons of the descriptive statistics for biogeographic variables among Diorhabda species are found in the species accounts below under the heading Biogeography - Comparative. Discussion of these results regarding biological control of tamarisk appear under the heading Potential in Tamarisk Biological Control for each species and in the concluding section, Implications Regarding Biological Control of Tamarisk

Key to the Sexes of the Diorhabda elongata Group: Adult External Characters

1. Distinct apical notch present on last visible abdominal sternite (Fig. 10). Third bi-lobed tarsomere of foretarsus 0.48–0.65 times the length of the fifth (final) tarsomere (Fig. 12) ...... male - Apical notch absent on last visible abdominal sternite (Fig. 11). Third bi-lobed tarsomere of foretarsus 0.28–0.44 times the length of the fifth tarsomere (Fig. 13) ...... female

Key to the Species of the Diorhabda elongata Group: Adult Males

1. Elongate (ventral) endophallic sclerite (EES) armed with spines on blade (SL) extending over an area less than or equal to 0.16 times (or less than about one fifth) the length of the sclerite, and blade (LB) extending less than or equal to 0.42 times the total length of the sclerite; EES never bearing a lateral appendage, lateral notch (pointed basally), or hooked apex (Figs. 14, 19, 24). Palmate (dorsal) endophallic sclerite (PES) with distal margin usually broadly rounded and with one to six (commonly two to four) usually subdistal spines (SDS) (maximum of two distal spines, DS), and no lateral appendage (rarely with a lateral notch, LN) (Figs. 14, 29). Subsutural (SSV) and submarginal elytral vittae (SMV), if present, never extending from apical half of elytra into the basal half (Fig. 1). Length 5.3–6.8 mm. Circummediterranean, Portugal, Bulgaria, Macedonia, S. Russia (Dagestan Republic), W. North America (introduced), commonly collected from coastal C. Turkey to Italy ...... elongata (Brullé) - Elongate endophallic sclerite armed with spines on blade extending over an area greater than or equal to 0.31 times (or greater than about one third) the length of the sclerite, with blade extending greater than or equal to 0.43 times the total length of the sclerite; EES sometimes bearing a lateral appendage (LA), lateral notch (LN, pointed basally) or hooked apex (HA) (Figs. 15–18, 20–23, 25–28). Palmate endophallic sclerite with distal margin truncate serrate and with two to six (commonly three to five) usually distal spines (maximum one spine subdistal) and a lateral appendage (Figs. 15–16, 30–31), or with distal margin narrowly or acutely rounded and one or two small subdistal spines and no lateral appendage (sometimes with lateral notch) (Figs. 17–18, 32–33). Subsutural and submarginal elytral vittae, if present, often extending from apical half of elytra into the basal half (Figs. 5, 9). Length 4.2–7.3 mm. Portugal, Spain, France, N. Africa, Senegal, Yemen, S. Ukraine, S. Russia, Turkey, Syria, Transcaucasus, Iraq, Iran, C. Asia, Pakistan, N.W. and N.C. China, S.W. Mongolia, W. North America (introduced)...... 2 2(1). Palmate endophallic sclerite bearing a strong lateral appendage and its distal margin truncate-serrate with two to six (commonly three to five) usually distal spines (maximum one spine subdistal) (Figs. 15–16, 30–31). Elongate endophallic sclerite with spines along blade often irregularly spaced with conspicuous gaps (Figs. 15–16, 20–21). Length 5.0–7.3 mm. Portugal, Spain, France, N. Africa, Senegal, Yemen, S. Ukraine, Turkey, Syria, Transcaucasus, C. Asia (east to N.W. China and Pakistan), Iraq, Iran, W. North America (introduced) ...... 3 - Palmate endophallic sclerite never bearing a lateral appendage (rarely bearing a lateral notch) and its distal margin narrowly or acutely rounded, with one or two small subdistal spines that may project beyond the distal margin (Figs. 17–18, 32–33). Elongate endophallic sclerite with spines usually evenly and closely spaced along blade (Figs. 17–18, 22–23). Length 4.2–6.1 mm. S. Russia, Transcaucasus, Syria, Iran, C. Asia, Pakistan, N.W. and N.C. China, S.W. Mongolia, W. North America (introduced) ...... 4 3(2). Lateral appendage (LA) of elongate endophallic sclerite always present and connected to lateral appendage of palmate endophallic sclerite by thin linear connecting (lateral) endophallic sclerite (CES) (Figs. 16, 21, 31) (in some weakly sclerotized specimens, the lateral appendage of the elongate endophallic sclerite and the connecting endophallic sclerite may be faint and appearing to evanesce). Length 5.0–6.9 mm. Portugal, Spain, France, N. Africa, Senegal, Yemen, Iraq, W. North America (introduced)...... sublineata (Lucas) - Lateral appendage of elongate endophallic sclerite usually absent or very weak and never connected to lateral appendage of palmate endophallic sclerite by a linear connecting endophallic sclerite (Figs. 15, 20, 30) (in some darkly sclerotized specimens, a faint line connecting the lateral appendages of the palmate and elongate

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 39 endophallic sclerites should not be interpreted as a developed connecting endophallic sclerite). Length 5.1–7.3 mm. S. Ukraine, Turkey, Syria, Transcaucasus, C. Asia (east to N.W. China and Pakistan), Iraq, Iran, W. North America (introduced)...... carinata (Faldermann) 4(2). Apex of elongate endophallic sclerite not hooked in dorsal aspect, but often bearing a lateral notch (Fig. 27; LN); blade of elongate endophallic sclerite extending from 0.48–0.66 times the total length of the sclerite (Figs. 17, 22; LB). Ratio of width to length of palmate sclerite ranging from 0.61–1.02; distal margin of palmate endophallic sclerite acutely rounded (Figs. 17, 32). Length 4.6–6.1 mm. S. Russia, Transcaucasus, Iran, C. Asia, N.W. and N.C. China, S.W. Mongolia, W. North America (introduced) ...... carinulata (Desbrochers) - Apex of elongate endophallic sclerite strongly hooked in dorsal aspect (Fig. 28; HA); blade of elongate endophallic sclerite extending from 0.67–0.84 times the total length of the sclerite (Figs. 18, 23). Ratio of width to length of palmate sclerite ranging from 0.35–0.63; distal margin of palmate endophallic sclerite narrowly rounded (Figs. 18, 33). Length 4.2–5.6 mm. Syria, Iran, Pakistan ...... meridionalis Berti & Rapilly

Key to the Species of the Diorhabda elongata Group: Adult Females

1. Vaginal palpi (VP) wider than long, with a width to length ratio (LP/WP) of 0.46–0.94 (Figs. 34–36). If width to length ratio of vaginal palpus 0.94, then the width of the widest lobe on the stalk (WLS) of internal sternite VIII (IS VIII) less than or equal to 0.10 mm (Figs. 34, 39). Length 4.9–8.4 mm. Circummediterranean, Senegal, Yemen, Portugal, Bulgaria, Macedonia, S. Ukraine, Turkey, Syria, Transcaucasus, C. Asia (east to Kazakhstan and Pakistan), Iraq, Iran, W. North America (introduced) ...... 2 - Vaginal palpi about as long as wide or longer, with a width to length ratio of 0.94–1.36 (Figs. 37, 38). If width to length ratio of vaginal palpus 0.94, then the width of the widest lobe on the stalk of internal sternite VIII greater than or equal to 0.11 mm (Figs. 37, 42). Length 4.8–7.0 mm. S. Russia, Transcaucasus, Syria, Iran, C. Asia, Pakistan, N.W. and N.C. China, S.W. Mongolia, W. North America (introduced) ...... 5 2(1). Vaginal palpi (VP) broadly rounded (Fig. 34). Width of the widest lobe on the stalk (WLS) of internal sternite VIII (IS VIII) from 0.06–0.11 mm (Fig. 34, 39). Subsutural (SSV) and submarginal elytral vittae (SMV), if present, never extending from apical half of elytra into the basal half (Fig. 1). Length 5.8–7.7 mm. Circummediterranean, Portugal, Bulgaria, Macedonia, S. Russia (Dagestan Republic), W. North America (introduced), commonly collected from coastal C. Turkey to Italy...... elongata (Brullé) - Vaginal palpi triangulate, narrowly rounded (Figs. 35–36). Width of the widest lobe on the stalk of internal sternite VIII from 0.08–0.18 mm (Figs. 35–36, 40–41). Subsutural and submarginal elytral vittae, if present, often extending from apical half of elytra into the basal half (Fig. 5). Length 4.9–8.4 mm. Portugal, Spain, France, N. Africa, Senegal, Yemen, S. Ukraine, Turkey, Syria, Transcaucasus, C. Asia (east to N.W. China and Pakistan), Iraq, Iran, W. North America (introduced) ...... 3 3(2). Stalk of internal sternite VIII (IS VIII) with tips of both lobes (TL) strongly curved inward away from apical lobe, and tips of lobes rounded or pointed, not quadrate (Fig. 40—Baghdad, Ashgabat, and Pul-e Charki). Length 5.1–8.4 mm. S. Ukraine, Turkey, Syria, Transcaucasus, C. Asia (east to N.W. China and Pakistan), Iraq, Iran, W. North America (introduced)...... carinata (Faldermann) - Stalk of internal sternite VIII with tip of at least one lobe not strongly curved inward away from apical lobe, and tips of lobes rounded, quadrate or pointed (Figs. 35–36, 40 – Lagodekhi and Ardanuc, 41 – , Tamri, Ndiol, and Perpignan) ...... 4 4(3). Stalk of internal sternite VIII (IS VIII) with tips of lobes (TL) either not curved or curved outward towards apical lobe, and tips of lobes rounded or quadrate, not pointed (Fig. 41- Ndiol, Perpignan, Kom Ombo). Length 4.9–7.4 mm. Portugal, Spain, France, N. Africa, Senegal, Yemen, Iraq, W. North America (introduced) ...... sublineata (Lucas) - Stalk of internal sternite VIII with tip of at least one lobe curved at least slightly inward away from apical lobe, and tips of lobes rounded, quadrate or pointed (Figs. 35–36, 40 – Lagodekhi and Ardanuc, 41 – Biskra and Tamri). Length 4.9–8.4 mm. (Iraq is area of range overlap) ...... carinata (Faldermann) and sublineata (Lucas) 5(1). Width of stalk (WST) of internal sternite VIII (IS VIII) 0.36–0.57 mm and this width 0.49–0.77 times the width of the apical lobe (WAL) (Figs. 37, 42). Length 5.0–7.0 mm. S. Russia, Transcaucasus, Iran, C. Asia, N.W. and N.C. China, S.W. Mongolia, W. North America (introduced) ...... carinulata (Desbrochers) - Width of stalk of internal sternite VIII 0.22–0.33 mm and this width 0.33–0.48 times the width of apical lobe (Figs. 38, 43). Length 4.8–6.4 mm. Syria, Iran, Pakistan ...... meridionalis Berti & Rapilly

40 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURES 1–9. Dorsal habitus of males (dead - top row, living - bottom row) of the Diorhabda elongata species group (tamarisk beetles): 1–2—D. elongata (Mediterranean tamarisk beetle), 3–4—D. carinata (larger tamarisk beetle), 5–6—D. sublineata (subtropical tamarisk beetle), 7–8—D. carinulata (northern tamarisk beetle), 9—D. meridionalis (southern tamarisk beetle). SSV—subsutural elytral vitta, SMV—submarginal elytral vitta. Scale bar 5.0 mm.

FIGURES 10–13. Sexual dimorphism in Diorhabda elongata: 10—male last visible sternite, 11—female last visible sternite, 12—male foretarsus, 13—female foretarsus. AN—apical notch, T3—third bi-lobed tarsal segment, T5—fifth tarsal segment. Scale bars 1.0 mm.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 41 FIGURES 14–18. Male genitalia: median lobe (ML) of aedeagus with everted endophallus (END; uninflated) (lateral and dorsal views).14—Diorhabda elongata, 15—D. carinata, 16—D. sublineata, 17—D. carinulata, 18—D. meridionalis. BF—basal foramen (or basal orifice), CES—connecting endophallic sclerite, DO—dorsal ostium (or apical orifice); EES—elongate endophallic sclerite, GP—gonopore, LA—lateral appendage, LB—length of blade, PES—palmate endophallic sclerite, SL—length of spined area of blade, VM—ventral membrane. Scale bar 1.0 mm.

42 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURES 19–23. Elongate (ventral) endophallic sclerite (lateral view) with connecting (lateral) endophallic sclerite (dorsal view, when present). 19—Diorhabda elongata, 20—D. carinata, 21—D. sublineata, 22—D. carinulata, 23—D. meridionalis. CES—connecting endophallic sclerite, EES—elongate endophallic sclerite, LA—lateral appendage, LB—length of blade, LN—lateral notch, SL—length of spined area of blade. Scale bar 1.0 mm.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 43 FIGURES 24–28. Elongate (ventral) endophallic sclerite (dorsal views and a single dorso-lateral view [DLV]). 24—Diorhabda elongata, 25—D. carinata, 26—D. sublineata, 27—D. carinulata, 28—D. meridionalis. BL—blade, HA —hooked apex, CES—connecting endophallic sclerite (partial), EES—elongate endophallic sclerite, LA—lateral appendage, LB—length blade, LN—lateral notch. Scale bar 1.0 mm.

Diorhabda elongata (Brullé, 1832) Mediterranean tamarisk beetle (Figs. 1, 2, 10–13, 14, 19, 24, 29, 34, 39, 44)

Galeruca elongata Brullé, 1832:271 (Type locality: Morée [Pelopónnisos peninsula, Greece]; Reiche and Saulcy, 1858:42 (part, France, Italy, Greece, Turkey, Syria, Lebanon [then under Syria], Egypt, Algeria, as Galleruca); Joan- nis, 1866:83 (part, Greece, Cilicia [Turkey], Lebanon [then under Syria], as Galleruca). Galeruca costalis Mulsant, Mulsant and Wachanru, 1852:176 (Type locality: Cilicia [in southwest Turkey], as Galle- ruca); Reiche and Saulcy, 1858:42 (established synonymy, as Galleruca); Wilcox, 1971:63 (world catalog). Diorhabda elongata: Weise, 1883:316 (part); 1893:635 (part; Italy, coastal env. Trieste), 1893:1132 (part, taxonomy), 1924:78 (part, world catalog, coastlands of Mediterranean Sea), 1925:225 (part, Egypt, area of Mediterranean Sea); Heyden et al., 1891:375 (part, catalog for Europe and Caucasus, southern Europe); Bedel, 1892:158 (part, France, as Dirrhabda); Winkler, 1924–1932 (part, Palearctic catalog, Mediterranean region); Corréa de Barros, 1924:9 (part, Portugal); Peyerimhoff, 1926:359 (part, Algeria, hosts); Laboissière, 1934:53 (part, France); Porta, 1934:317 (taxo- nomic keys, Italy); Ogloblin, 1936:79 (part, Italy [Trieste], Greece, Turkey, Syria, Lebanon [Palestine], S. Russia); Kerville, 1939:107 (Turkey, env. Smyrna); Hopkins and Carruth, 1954:1129 (part, Spain, host); Pavlovskii and Shtakelberg, 1955:566 (part, Mediterranean region, southwest Russia, as Diorrhabda); Torres Sala, 1962:327 (part, Comunidad Valenciana, Spain); Lopatin, 1967:441 (Lebanon, dunes south of Beirut); Tomov, 1969:181 (Bulgaria), 1979:165 (biology, Bulgaria), 1984:377 (part, Turkey); Jolivet, 1967:331 (part, Mediterranean, hosts); Zocchi, 1971:86 (part, Italy); Wilcox, 1971:63 (part, world catalog); Gerling and Kugler, 1973:20 (Israel [sic; should read Turkey]); Warchalowski, 1974:509 (Bulgaria, as Diorrhabda), 2003:328 (part, taxonomic keys, Mediterranean region, Caucasus); Tomov and Gruev, 1975:146 (Turkey); Georghiou, 1977:47 (Cyprus); Lundberg et al., 1987b:126 (Sicily); Petitpierre, 1988:93 (part, Spain); Regalin, 1997:69 (Crete); Gruev and Tomov, 1986:103 (Bulgaria), 1998:70 (part, Bulgaria, Mediterranean); Biondi et al., 1995:12 (Italy); Kovalev, 1995:78 (part, southwest Palaearc- tic); Campobasso et al., 1999:145 (part, host, Europe and Middle East); Lair and Eberts, 2001:1 (introduction plans, north Texas); Aslan et al. 2000:30 (part, Turkey); Anonymous, 2001:52N (part, southern Europe); Chatenet, 2002:223 (part, Italy, France, Spain, Algeria); DeLoach et al., 2003a:229 (part, Greece), 2003b:126 (part, host range,

44 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS ecology, Italy, Crete, Turkey), (2008, in prep.) (part, Greece); Gök and Çilbiroğlu 2003:66 (Turkey), 2005:14 (Tur- key); Milbrath et al., 2003:225 (part, Greece); Riley et al., 2003:69,189 (part, catalog North America [introduced]); Bieńkowski, 2004:76 (part, keys, eastern Europe [Southern Russia]); DeLoach and Carruthers, 2004a:13, 2004b:311 (part, Greece); Gök and Duran, 2004:17 (part, Turkey); Lopatin et al., 2004:127 (part, Mediterranean); Dudley, 2005a:13, 2005b:42N (part, biological control, ex: Greece); Carruthers et al. 2006:71, 2008:262 (part, biological control, ex: Greece); Herr et al. 2006:148 (host specificity, Crete); Milbrath and DeLoach, 2006a:32, 2006b:1379 (part, host specificity, Crete); Milbrath et al., 2007 (part, host specificity, biology, Crete); Dudley et al., 2006:137 (releases in California, Crete); Everitt et al. (2007) (remote sensing, Texas, Crete); Hudgeons et al., 2007a:157 (establishment in Texas, Crete), 2007b:215 (tamarisk damage in Texas, ex: Crete); Mityaev and Jashenko, 2007:145 (biological control, ex: Greece); DeLoach (2008); Moran et al. (in press) (host range in Texas, Crete); Bean and Keller (in prep.) (diapause induction, Crete); Dalin et al. (in press) (host range; Crete); Thompson et al. (in prep.) (part, laboratory hybridization, Crete). Diorhabda elongata ab. carinata: Porta, 1934:317 (keys, Italy).

Male. Genitalia. Diorhabda elongata can be distinguished from all other members of the D. elongata group by the following combination of characters, the first two of which are unique among the tamarisk beetles: (1) the length of the spined area of the blade (SL) (armed with one to six [commonly one to three] spines) of the elongate endophallic sclerite (EES) is less than or equal to 0.16 times (or less than about one fifth) the length of the EES (Table 3; Figs. 14, 19, 48B); (2) the length of the blade of the EES (LB) is less than or equal to 0.42 times the length of the EES (Table 3; Figs. 14, 19, 24, 48A); (3) the EES lacks a lateral notch (pointed basally), lateral appendage or hooked apex (Figs. 14, 19); (4) the palmate endophallic sclerite (PES) lacks a lateral appendage, it is usually broadly rounded distally, and the one to six (commonly two to four) spines are usually subdistal with no more than two distal spines (Figs. 14, 29); and (5) the connecting endophallic sclerite is lacking. In the other four species of the D. elongata group, the length of the spined area of the EES (SL) (armed with three to seven spines) is greater than or equal to 0.31 times (or greater than about one third) the length of the EES (Table 3; Figs. 20–23, 48B), and the length of the blade of the EES is greater than or equal to 0.43 times the length of the EES (Table 3; Figs. 15–18, 20–23, 25–28, 48A). In D. sublineata,and sometimes D. carinata, the EES bears a lateral notch (pointed basally) or lateral appendage. In D. meridionalis the EES bears a hooked apex (Figs. 25–28). The PES always bears a lateral appendage and the distal margin is truncate serrate with usually more than two distal spines in D. carinata and D. sublineata (Figs. 15–16, 30–31). In D. carinulata and D. meridionalis, the distal margin of the PES differs from that of D. elongata in being narrowly or acutely rounded with one or two small subdistal spines (Figs.17–18, 32–33). Measurements. See Tables 2 and 3. Female. Genitalia. Female D. elongata may be distinguished from all other members of the D. elongata group by the following combination of characters in the vaginal palpi (VP) and internal sternite VIII (IS VIII): (1) the vaginal palpi are both broadly rounded and wider than long with a width to length ratio (LP/WP) of 0.52–0.94 (Fig. 34, Table 4), and (2) if the width to length ratio of the vaginal palpi is 0.94, then the width of the widest lobe of the stalk (WLS) of IS VIII is less than or equal to 0.10 mm (Fig. 34). In contrast to D. elongata, the vaginal palpi in D. carinulata (Fig. 37) and D. meridionalis (Fig. 38) are about as long as wide or longer with a width to length ratio of 0.94–1.36 (Table 4). If the width to length ratio of the vaginal palpus is 0.94 in D. carinulata, then the width of the widest lobe of IS VIII is greater than or equal to 0.11 mm (Fig. 37). The vaginal palpi of D. carinata and D. sublineata differ in being narrowly rounded and triangulate (Figs. 35–36). In addition, the width of the widest lobe of the stalk of IS VIII is generally smaller in D. elongata (range 0.06–0.11 mm; Fig. 34) compared to D. carinata (range 0.11–0.17; Fig. 35) and D. sublineata (range 0.08–0.18 mm; Fig. 36) (Table 4). Measurements. See Tables 2 and 4. Coloration. Diorhabda elongata commonly lacks elytral vittae, but submarginal and subsutural vittae in the apical half of the elytra (evanescing toward the basal half) are not uncommon (Fig. 1). In all other members of the D. elongata group, elytral vittae, when present, often extend into the basal half of the elytra (Figs. 5, 9). Live specimens of D. elongata possess greenish-yellow tinting along veins of the elytra, probably as a result of yellow hemolymph as seen in live inflated endophalli, and this gives adults an overall olivaceous

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 45 hue (Fig. 2). Living D. carinata have a lesser degree of greenish-yellow tinting (Fig. 4). Living D. sublineata (Fig. 6) and D. carinulata (Fig. 8) lack greenish-yellow tinting, probably as a result of white hemolymph seen in live inflated endophalli (also seen in D. carinata), and they appear more tannish-yellow in hue. Type material. Brullé’s collection from his expedition to Morée (= Pelopónnisos peninsula, Greece), which should include the type specimen(s) for D. elongata, was deposited in the Muséum National d'Histoire Naturelle in , France (MNHN) (Groll 2006). A curator at MNHN communicated intent to inform us of the status of the type material and perhaps lend it for examination, but after four years, we have not heard of the status of the Brullé type material. Once it can be ascertained that the type material is lost, a neotype should be designated using a dissected male specimen from the type locality of Pelopónnisos, Greece. We studied the original description by Brullé (1832), based on an unspecified number of specimens, with the included color habitus illustration, and topotypes from the Pelopónnisos peninsula, Greece. The location of type specimens of Galeruca costalis Mulsant (Mulsant and Wachanru 1852) is uncertain. Remaining portions of E. Mulsant’s collection “Natural History of the Coleoptera of France” should be located at the l’Institution Sainte–Marie de Saint–Chamond (Loire), France (Groll 2006). We studied the original descriptions of G. co s ta li s by Mulsant (Mulsant and Wachanru 1852) and topotypes from the Cilicia region of Turkey. Material examined. 157♂♂ dissected (diss.), 85♀♀ diss., 127♂♂, 164♀♀. ALBANIA: 1♂ diss., Durazzo [41.32305°N, 19.44138°E], Lona-Rav., Diorhabda elongata det. Burlini 1967, MSNM [2003-08]; 1♂ diss., 1♂, 1♀, Elbasan [41.11250°N, 20.08222°E], D. elongata, USNM [2003-10]; 1♂ diss., Medua [Shengjin, San Giovanni di Medva; 41.81366°N, 19.5939°E], Matzenaver, D. elongata det. V. Zonfal, NMPC [2004-51]; 1♂ diss., Srutari [Shkoder; 42.0675°N, 19.5131°E], D. elongata det. Burlini 1967, MSNM [2003- 11]; 1♀ diss., Scutari [Shkoder], Heyrevsky, NMPC [2006-06]; 1♂ diss., 1♀, Valona [Vlore; 40.46666°N, 19.48972°E], [19]08, H. Hopp, ZHMB [2003-04]; ALGERIA: 1♂ diss., [specific locality not given], Reitter, NMPC [2004-15]; BOSNIA & HERZEGOVINA: 1♂ diss., Veleś Planina [Velez Planina; 43.33°N, 18.0044°E], Herzegowina, 1879, Reitter, D. elongata, HNHM [2003-05]; BULGARIA: 1♂ diss., 1♂, 1♀, [42.5°N, 27.4667°E], 16-VII-1975, Vaśárhelyi, D. elongata det. V. Tomov 1983, HNHM [2003-06]; 1♀ diss., Gara [Kresna; 41.7333°N, 23.1500°E], Kresn. defile [Kresnenska Klisura or Kresna defile (gorge)], B. Kouřil, Diorhabda det. B. Kouřil, P9/46/62, NMPC [2005-25]; 1♂ diss., 1♀, Gara [Sandanski; 41.5667°N, 23.2833°E], [river], 20-VII-1956, L. Hoberlandt, NMPC [2006-04]; 1♂ diss., Kresna [41.7333°N, 23.1500°E], environs, Struma [river], 29-V-1984, U. Göllner, D. elongata det. V. Tomov, ZMHB [2003-10]; 1♂ diss., 1♂, 1♀, Kresnicko Def. [Kresnenska Klisura or Kresna defile (gorge); 41.8°N, 23.16667°E], VII-1932, Mac, Mař et Táb, NMPC [2006-05]; 1♀ diss., Kritschin [; 42.05°N, 24.4667°E], NHMB [2003-41]; 1♂ diss., Melnik [41.5167°E, 23.4°E], 23-V-1973, St. Gruszka & A. Warchalowski, GSWRL [2003-41]; 2♂♂ diss., 1♀diss., Petric [42.6°N, 24.01667°E], 1932, D. Putkyně, D. elongata det. Sterba, NMPC [2004-24, 2005-11, 13]; 1♀ diss., [42.55°N, 27.65°E], 9-V-1970, B. Gruev, D. elongata det. V. Tomov 1997, GSWRL [2003-46]; 1♂ diss., 3♂♂, 2♀♀, Sandanski [41.5667°N, 23.2833°E], Meridion., 10-VIII-1974, A. Warchalowski, GSWRL [2003-34, 1♂ diss.], DEI; 1♂ diss., 4♂♂, 4♀♀, Keretschkoi [Kirechdzhi Khask'oy is Sladun; 41.85°N, 26.4667°E], Macedonia, A. Schatzmayer, D. elongata det. Fleischer, DEI [2003-18]; CROATIA: 1♂ diss., Metkovic [Metkovi; 43.054167°N, 17.648333°E], Dalmatien, E.A. Bottcher (Berlin), 168, MZHF [2005-03]; 1♂ diss., Metkovich [Metkovi], Dalmatia, NHMB [2003-25]; 1♀ diss., 1♀, Opuzen [43.00972°N, 17.56555°E], 26-V-1974, G.J. Slob, RBCN [2003-07], ZMAN; 1♂ diss., 5♂♂, 10♀♀, Ušce Neretve D. [43.01917°N, 17.44389°E; Neretva river mouth], 8-IX-1948, Novak [ZMHB], D. elongata det. Novak, HNHM [2003-04], ZMHB [2♂♂, 4♀♀]; CYPRUS: 1♂ diss., [specific locality not given], 1876, L. Shrader, Coll. Kunnemann, DEI [2003-13]; 2♂♂ diss., 1♀ diss., 6♂♂, 5♀♀, Larnaka [Larnaca; 34.916667°N, 33.6333°E], 25-VI to 1-VII-1939, Håkan Lindb., D. elongata det. Harald Lindb. [3♂♂, NMPC; 1♂, 1♀, NHRS], MZHF [2003-02, 2005-01, 02], NMPC, NHRS; 1♂, 1♀, Larnaca, Kamares Aqueduct, 19-X-1932, A. Ball, D. elongata det. V. Laboissière 1939, IRSNB; 2♂♂ diss., 1♀ diss., 3♀♀, Mt. Arménien [35.2875°N, 33.525°E], NMPC [2004-02, 2005-01, 02]; EGYPT: 3♂♂ diss., 2♀♀ diss., 1♀, Cairo [Al Qahirah; 30.05°N, 31.25°E], NHMB [2003-16, 52, 2004-01a, 2005-02, 03];

46 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS GREECE: 1♂ diss., Agia Marina, Star Beach [35.5193°N, 23.9275°E], Crete, 10-IX-2001, R. & N. Carruthers, on Tamarix sp. foliage, EIWRU-2001-1016, USNM [2003-18]; 1♀ diss., 3♀♀, Ammos [36.91270°N, 27.28170°E], Kos [Island], 27-VIII to 1-IX 1981, T. Palm, MZLU [2005-08]; 6♂♂ diss., 2♀♀ diss., Astro Beach [37.4433°N, 22.74861°E], north of, Pelopónnisos, 25-V-2003, J. Kashefi, on T. hampeana, GSWRL [2004-10, 2005-38, 39, 41–45]; 1♀diss., Athens environs [37.96138°N, 23.63888°E], Dr. Krüper, NHMB [2004-03]; 1♂, 2♀♀, b. Athens, v. O., Collectie C. et O. Vogt Acq. 1960, ZMAN; 1♂ diss., 1♂, 2♀♀, Athos [40.166667°N, 24.333333°E], Macedonia, Schatzmayer, Coll. V. Heyden, D. elongata det. Fleischer, DEI [2003-19]; 2♂♂ diss., 3♂♂, Athos, Macedonien, 1908, Schatzm., NHMB [2003-26, 2006-02]; 1♀diss., 1♂, 4♀♀, Attica [Attiki; 38.083333°N, 23.5°E], Stussiner, O. Leonard, DEI [2003-15]; 1♂, 3♀♀, Attica, Reitter, Coll[ection] Kambersky, D. elongata, NMPC; 2♂♂ diss., 1♂, 2♀♀, Attica, Reitter, 269 [MZLU ♀] D. elongata Brulle Emm. Reitter [ZMAN; 1♂, 1♀], Collectie P.v.d. Wiel Acq. 1962 [ZMAN], MZLU, ZMAN [2008-05, 17], ZMHB [1♂]; 1♂, Attica, Witte, coll[ection] V. Heyden, DEI; 1♀, Attica, Hymettos [Mts.], Emge., ZMAN; 1♀, Attica, v. Oertzen, ZMAN; 1♂, Attika, ZMHB; 1♀ diss., 1♂, 1♀, Crete [specific locality not given], D. elongata det. K. Lopatin, HNHM [2005-01]; 1♀diss., 1♀, Diakophto [Diakofto; 38.1795°N, 22.2001°E], Pelop., IV-1936, Mař et Táb., Coll[ection] Bartoň, NMPC [2004-34]; 1♀ diss., Dimilia [36.2834°N, 28.0085°E], Rhodes [island], 16-VII-1933, A. Mochi, D. elongata det. Burlini 1967, MSNM [2003-14]; 1♂ diss., Euboea [Evvia Island; specific locality not given; approx. loc.: 38.5200°N, 23.72000°E], Coll. Kraatz, DEI [2003-14]; 1♂ diss., 1♀, Fileremo [Filerimos Hill; 36.3985°N, 28.1382°E], Rodi [Rhodes], 22-IV-1932, A. Schatzmayer, D. elongata det. Burlini 1967, MSNM [2003-12]; 1♀ diss., Fileremo [Filerimos Hill], Rodi [Rhodes], 10-VII-1933, A. Mochi, D. elongata det. Burlini 1967, MSNM [2008-02]; 1♂ diss., Heraklion [Irakleion; 35.325°N, 25.1306°E], Crete, 13-IX-2001, R. & N. Carruthers, lab reared from eggs collected from Tamarix sp. foliage, EIWRU-2001-1016, USNM [2003-09]; 1♀ diss., 1♀, Heraklion [Irakleion], 18-X-2001, R & N. Carruthers, lab reared from eggs on Tamarix spp. [at Albany, CA], USNM [2005-01]; 1♀ diss., 1♂, 1♀, Heraklyon [Irakleion], Kreta, 22-V-1995, J. Blümel, D. elongata det. Erber 2001, ZMHB [2008-04]; 1♀ diss., 2♂♂, 1♀, Hypati [Ypati; 38.88590°N, 22.23350°E], Pfeffer, D. elongata det. V. Tomov 1984 [1♂ diss.], NMPC [2004-46]; 1♂ diss., 1♂, 3♀♀, Hypati [Ypati], IV-1936, Mař et Táb., Coll[ection] Bartoň, NMPC [2006-07]; 1♂ diss., 6♂♂, 3♀♀, Ilis [37.8833°N, 21.3833°E], Olympia, 2–3-X- 1962, Ent. Exc.Zoöl Mus., ZMAN [2008-14]; 1♂ diss., 2♂♂, 3♀♀, Ixia [Ixos; 36.4167°N, 28.18330°E], Rhodes, 19-IV to 1-V-1973, A. Teunissen, RBCN [2003-04], ZMAN [1♀]; 1♂ diss., Kalamata [37.0389°N, 22.1142°E], 22-IX-1982, Roland Bö, 579, D. elongata det. Eber 1989, ZMHB [2008-01]; 1♂ diss., Karia environs [39.98330°N, 22.4000°E], Olymp[us Mt.], 1000 m [elev.], 30-VI-1980, Probdt, D. elongata det. V. Tomov 1983, NMPC [2005-26]; 1♂ diss., 4♂♂ Kardamena [Kardamaina; 36.7814°N, 27.1425°E], Kos [Island], 2–6-IX-1981, T. Palm, MZLU [2005-01]; 2♀♀ diss., 1♂, 5♀♀, Karpathos Town [35.5°N, 27.23333°E], Ormos Pigadia env., 0–150 m, Karpathos Island, 28-VI to 3-VII-1996, W.T. Edzee, D. elongata det. A. Teunissen 1998 [ZMAN ♀], RBCN [2003-12, 2005-01; 2♀♀], ZMAN; 1♀diss., Katakolo [Katakolon; 37.6500°N, 21.3167°E], Morea v. O., Collectie C. et O. Vogt Acq. 1960, ZMAN [2008-15]; 1♂ diss., 2♂♂, Kerkyra [39.6200°N, 19.91970°E], Korfu [Kérkyra Island], V-1964, Palm, MZLU [2005-07]; 1♂ diss., 1♀, Katafourko [Katafourkon; 38.9981°N, 21.1511°E], (Etol), 27–30-V-1998, P. Foot, ZMAN [2008-04]; 1♀diss., 1♂, 1♀, Korfu, Insel [Kérkyra Island; specific locality not given], NHMB [2003-28] [1♂ resembles aberration bipustulata]; 1♂ diss., 6♀♀, Korission [Korissíon, Limni (sea); 39.44611°N, 19.90694°E], Korfu [Kérkyra Island], V-1964, Palm, MZLU [2005-06]; 1♂ diss., 1♀, Kos [Island, specific locality not given], Südl. Sporaden, Oertzen, D. elongata, 86202, ZMHB [2003-11]; 2♂♂ diss., 1♀ diss., Kyparrisia [37.2577°N, 22.6533°E], Pelopónnisos, 26-V-2003, J. Kashefi, on T. parviflora, GSWRL [2004-11, 23, 24]; 1♂ diss., Lixourion [38.201944°N, 20.431389°E], Cephalonia, 26-VIII-1899, A. Porta, D. elongata det. A. Porta, MSNM [2003-15]; 1♀ diss., Mesolongion [38.369167°N, 21.429167°E], NHMB [2003-40]; 1♂ diss., Malia Beach [Mália; 35.2883°N, 25.4667°E], meadow west of hotel, Kreta [Crete island], 5–12-V-1979, R. Danielsson, D. elongata det. Mohr 1980, MZLU [2005-12]; 4♂♂ diss., 2♀♀ diss., 3♂♂, 4♀♀, Methoni [36.81666°N, 21.7°E], Peloponneso [Pelopónnisos Penninsula], 23-IX-1997, MRSN [2003-02, 09, 10, 11, 12, 13]; 3♂♂, 1♀, Morea [Pelopónnisos Penninsula], v. Oertzen (2♂♂, 1♀), Collectie C. et O. Vogt Acq. 1960,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 47 ZMAN; 1♂ diss., 1♂, Oeta [Mt.; specific locality not given], IV-1936, Mař et Táb., Coll[ection] Bartoň, NMPC [2004-36]; 1♂ diss., 1♂, 1♀, Molyvos [Mithymna; 39.3667°N, 26.1667°E], Lesbos [Lesvos Island], 4–11-V-1992, J.H. Woudstra, ZMAN [2008-06]; 1♂ diss., 5♂♂, 6♀♀, Olympia [Archaía Olympia; 37.6500°N, 21.6333°E], Alpheios [Alfeiós] Valley, Morea [Pelopónnisos Penninsula], 6-V-1930, A. d'Orchymont, D. elongata det V. Laboissière 1939, IRSNB [2006-04]; 1♂ diss., Olympia [Archaía Olympia], Morea, v. Oertzen, ZMAN [2008-19]; 1♀ diss., 1♀, Pahia Ammos [Pakhia Ammos; 35.11670°N, 25.8000°E], Kreta [Crete Island], 16-V-1975, MZLU [2005-11]; 1♀ diss., 1♂, Pantokrator [Pantokrátor Mt.; 39.75396°N, 19.86277°E], Korfu [Kérkyra Island], V-1964, Palm, MZLU [2005-09]; 1♂ diss., 2♀♀, Paradisi [36.4006°N, 28.0796°E], Rhodes, 21-IV-1996, A. Teunissen, RBCN [2003-18], ZMAN [1♀]; 1♀ diss., Peloponn. [Pelopónnisos Penninsula], D. e. var. sublineata det. K. Lopatin 1958, HNHM [2004-06]; 1♂ diss., 1♀, Piraus [Peiraiefs] env., Athens, 2-VIII-1982, Steinhausen, D. elongata det. Steinhausen, RBCN [2003-14]; 1♀ diss., Pirgos [37.6833°N, 21.45°E], Peloponnese [Pelopónnisos Penninsula], 17-VI-1986, H. Hebauer, D. elongata det. Doberl 1989, MBPF [2003-05]; 1♀ diss., 1♂, 1♀, Plimmiri [35.929°N, 27.8584°E], Rhodes, 20-IV-1996, A. Teunissen, RBCN [2003-20], ZMAN [1♂]; 1♂ diss., 1♀ diss., Possidi [Posidi Beach; 39.96467°N, 23.36483°E], Kassandra, Halkidiki, 3-VI-1989, R. Sobhian, on Tamarix sp., “RS,89,I09”, USNM [2005-03, 04]; 3♂♂ diss., 2♀♀ diss., 27♂♂, 14♀♀, Possidi [Posidi Beach], Kassandra, 39° 57.88' N, 23° 21.89' E (GPS), 28-IV-1999, J. Kashefi & R. Sobhian, on Tamarix spp., 9063, 9091, 9092, 9094, 9095, 9097, 9098–9118, 9120–9124, D. elongata det. I.K. Lopatin 1999 [2♂♂ diss., 2♀♀ diss., 23♂♂, 8♀♀], GSWRL [2003-13, 2005-27, 28, 29, 2006-03]; 6♂♂ diss., 4♀♀ diss., Posidi Beach [not Thessaloniki, as in some shipping records; Halkidiki region], 3-X-2002, J. Kashefi, EIWRU-2002-1009, USDA/ARS lab colony at Temple, Texas, voucher [J.L. Tracy, 2002–2003], USDA/ARS lab colony at Albany, California, voucher, GSWRL [2003-17, 19, 26, 27, 30, 2005-33, 34, 35, 64, 65] [introduced in New Mexico and Texas in 2004] [used in biological studies Herr et al. (in prep.), and Bean and Keller (in prep.)]; 1♂ diss., Rhodos umg. [env.] [Rodos; 36.4408°N, 28.2225°E], 1-VIII-1982, Steinh., D. elongata R. Beenen det. 1996, ZMHB [2008-02]; 1♀, Rhodes [island, specific locality not given], Reitter, NMPC; 1♂ diss., 1♀, Rhodos [Rhodes island,], 15- VIII-1970, Rich Dahl, D. elongata det. L. Borowiec, MZLU [2005-05]; 1♀ diss., 1♀, Rhodos [Rhodes island], 24-VI to 4-VII-1958, Palm, MZLU [2005-10]; 1♀, Rhodes, Reitter, NMPC; 1♀, Salonich [Thessaloniki; 40.64027°N, 22.94388°E], Schatzmayer, coll[ection] V. Heyden, DEI; 1♂ diss., 1♀, Saloniki [Thessaloniki], Vardarebene [Vadar River], ZMHB [2003-02]; 1♂ diss., Servia, 5 km NW [40.2204°N, 21.9571°E], Makedonia, 19-VIII-1965, E. A. Blommers, ZMAN [2008-09]; 1♂ diss., 1♂, Sigri [Sigrion; 39.2167°N, 25.8500°E], Lesbos [Lesvos Island], 8-V-1992, J.H. Woudstra, ZMAN [2008-10]; 7♂♂ diss., 6♀♀ diss., 2♂♂, 3♀♀, Sfakaki, 3 km west [35.3833°N, 24.6°E; near Stavramenos, Crete], road from Irakleion to Rethymnon, 5-IV-2002, R. Carruthers & J. Kashefi, shipment EIWRU-2002-1002, USDA lab colony at Temple, Texas, voucher (28-VI-2002, 6-IX-2002, J.L. Tracy; 1♂ diss., 28-VIII-2002, 1♂, 2♀♀, 26- VIII-2002, T.O. Robbins; 1♂ diss., 1♀ diss, 30-VI-2003, 28-VII-2003, L. Milbrath [nos. 1278, 1274]), GSWRL [2003-08, 10, 20, 24, 25, 31, 32, 40, 46, 47, 79, 82, 83] [introduced in California, New Mexico and Texas in 2003] [used in biological studies of Milbrath and DeLoach (2006a, 2006b); Milbrath et al. (2007); Moran et al. (in press); Herr et al. (2006, in prep.); Bean and Keller (in prep.); Hudgeons et al. (2007a, 2007b); and Thompson et al. (in prep.)]; 1♀ diss., 3♀♀, Sparta [Sparti; 37.0733°N, 22.4297°E], Pelopones, 1935, Mařan et Stěp., Coll[ection] Bartoň, NMPC [2004-32]; 1♂ diss., Taygetos [Mt.; specific locality not given], Pelopon. [Pelopónnisos Penninsula], 1935, Dr. Purkyne, NMPC [2004-37]; 1♀ diss., Sparti, 5 km SW [37.0386°N, 22.3985°E], Lakonia [Laconia Prefecture], 27-IX-1962, Ent. Exe. Zoö. Mus., ZMAN [2008-08]; 1♀ diss., Thérisos [Therissos; 35.3333°N, 25.1167°E], 3 km W Iráklion [Irakleion], Kriti [Crete], 14-X-1972, A.C. & W.N. Ellis, ZMAN [2008-07]; 4♂♂ diss., 2♀♀ diss., 1♂, 9♀♀, Thessaloniki, 60 km E [40.64°N, 23.48°E], 117 m [elev.], 30-IV-1999, Kashefi & Sobhian, on Tamarix spp., 9126–9129, 9131–9136, D. elongata det I. Lopatin 1999 [5♂♂, 5♀♀], GSWRL [2003-40, 2005-26, 30, 51, 2006-04, 05]; 1♀ diss., Trianta [36.4°N, 28.16666°E], Rhodes, 29-IV-1932, A. Schatzmayer, D. elongata det. Burlini 1967, MSNM [2003-13]; 1♀ diss., Xylokastron [38.08333°N, 22.63333°E], 29-V-1964, E. Junger, ZMHB [2003-03]; ITALY: 1♂ diss., 1♀ diss., 2♂♂, Alcantara River banks by sea [37.8067°N, 15.2594°E], Sicilia [Sicily], V-

48 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS 1926, NHMB [2003-49, 2005-01]; 1♂ diss., 1♂, Ancona [43.6333°N, 13.5000°E], Zeller, 611, 172, DEI [2008-01]; 1♂ diss., 2♂♂, 3♀♀, Cotrone [Crotone, resembles Botrone; 39.0833°N, 17.1333°E], Calabria, 26- VIII-[18]88, A. Fiori, ZMHB [2008-06]; 1♂ diss., Fiumi di Pollina [38.01667°N, 14.16667°E], Sicilien [Sicily], 1982, T. Palm, D. elongata det. Thure Palm, coll[ection] T. Palm, MZLU [2005-02]; 1♀, Lido [45.6783°N, 13.4025°E], O. Funk, ex. coll. J. Weise, ZMHB; 1♂ diss., Lido di Volana [Lido di Volano; 44.79638°N, 12.27083°E], Littorale Ferrarese, 24-V-1959, A. Giordani Soika, Tamarix dunes, MRSN [2003- 07]; 1♂ diss., 1♂, Messina [38.1833°N, 15.5667°E], Sicilia [Sicily], Reitter, D. elongata [NMPC], Coll. Dr. J. Erdo’s [HNHM], HNHM [2004-07], NMPC; 1♂ diss., Messina, Fobria [?], 21-IX-1952, A. Porta, Galerucella grisescens det. Antonio Porto, MSNM [2008-01]; 4♂♂ diss., 5♀♀ diss., 2♂♂, 4♀♀, Monte Gargano [Promontorio de Gargano; 41.8333°N, 16.0°E], 30-IV to 13-V-1907, M. Hilf, Coll. O. Leonard, D. elongata [1♂], DEI [2003-01, 02; 2005-02, 03, 04], NMPC [2006-10 ♂; 2 ♀♀], ZMHB [2♀♀]; 1♂ diss., Pescara [42.4667°N, 14.2167°E], Abruzzo, 12-IV-1906, A. Fiori, ZMHB [2003-05]; 1♂ diss., 1♀, Piano Torre [resort near Altavilla Milicia, Sicily; 38.0333°N, 13.5333°E], X-1908, [ZIN 2004-12]; 1♀ diss., Pineta [42.45°N, 14.2333°E], Lido Bruno, Puglia Prov., 7-III-1981, F. Montemurro, seaside, D. elongata det. D. Sassi 1995, GSWRL [2003-60]; 1♂ diss., 1♀, 1♂, Pioppi ([nr.] Salerno) [40.1833°N, 15.0833°E], V–VI- 1965, H.K. Mohr, DEI [2003-09]; 1♀ diss., Pioppi ([nr] Salerno), 23-X-1964, W. Liebmann, DEI [2003-24]; 1♂ diss., 1♂, Rendina Val[ley] [Rendina River; approximate location: 41.0166°N, 15.7382°E], Leone [?], Basilicata, 1901, A. Fiori, ZMHB [2008-07]; 1♂ diss., Rodia [Villagio Rodia, suburb of Messina], X-1932, F. Vitale, Le Moult vend. via Reinbek Eing. 1-1957, 78, D. e. ab. carinata det. V. Laboissière, ZMUH [2006-01]; 1♂ diss., Rosalina Mare [45.1192°, 12.3133°E], 4-VII-1962, A.G. Soika, beach dune with Ammophiletum, MRSN [2003-05]; 1♀ diss., Sinnaro [Contrada de Sinnaro, quarter of Messina], 16-VI-1929, F. Vitale, Le Moult vend. via Reinbek Eing. 1-1957, 78, D. e. ab. sublineata det. V. Laboissière, ZMUH [2006-02]; 1♂ diss., 2♂♂, 2♀♀, Taormina env. [37.85°N, 15.283333°E], 8-V-1942, Frey, NHMB [2003-20]; 1♂ diss., Toscani [Toscana Province; specific locality not given], Tamarix gallica, Coll[ection] Chapuis, D. elongata det. V. Laboissière 1939, IRSNB [2006-09]; LEBANON: 1♀ diss., Beirut [33.87194°N, 35.50972°E], dunes south of, 10–15-V-1963, Kasy & Vartian, USNM [2003-20] [from series listed as D. elongata by Lopatin (1967)]; 4♂♂ diss., 1♀♀ diss., Beyrut [Beirut], Syr, Krüper, NMHB [2003-12, 13, 17, 45, 48]; 1♀ diss., Beyrut [Beirut], Syr, 20-IV-1936, Frey, NHMB [2003-38]; 2♂♂ diss., 1♀ diss., 1♀, Beyrut [Beirut], Syrien, NHMB [2003-46, 2005-04], ZMHB [2003-09]; MACEDONIA: 2♀♀, Macedonien [specific locality not given], Emge., ZMAN; 1♂ diss., Mazedonien [specific locality not given], 4-VI-1987, J. Böhme, 533, D. elongata det. Erber 1985, ZMHB [2008-05]; 1♂ diss., 2♀♀, 1♂, Skopje env. [42.0°N, 21.4333°E], southern Yugoslavia, VI-1937, O. Kodyn, D. elongata det. Sterbud, NMPC [2004-52]; 1♂ diss., 1♀ diss., 5♂♂, 3♀♀, Strumica [41.4375°N, 22.64333°E], 19-V-1937, W. Liebmann, DEI [2003-20, 2005-01]; 1♀ diss., Vardar [42.00555°N, 21.32611°E; stream], D. elongata det. Burlini 1967, MSNM [2003-09]; : 1♂ diss., Sutomore [42.14277°N, 19.04666E], 22-V-1978, G.J. Slob, RBCN [2003-03]; PORTUGAL: 2♂♂ diss., [specific locality not given], Coll. Letzner, DEI [2003-10, 2006-01]; RUSSIA: 1♂ diss., Daghestan [Respublika Dagestan; specific locality not given], Leder. Reitter, 4358, Coll[ection] Kouril P5/46/62, NMPC [2005-40]; SPAIN: 2♀♀ diss., 1♀, Estepona [36.4333°N, -5.1333°W], Malaga, 15-II-1982, H. Teunissen, RBCN [2003-13, 16]; SYRIA: 1♂ diss., [specific locality not given], D. e. var. carinata, MNMS [2004-04]; 1♀ diss, [specific localilty not given, D. elongata det. Le Moult, IRSNB [2006-11]; TURKEY: 1♂ diss., 1♀ diss., Adana [37.001667°N, 35.328889°E], D. elongata [1♂ diss., NHMB], HNHM [2005-02], NHMB [2003- 10]; 3♂♂ diss., 1♀ diss., 3♀♀, Adana, Asia Minor, Sterba, D. v. carinata det. TJS [1♀ undiss.], NMPC [2005-08–10, 12]; 1♀, Adana, Asia Minor, H. Rolle, ZMHB; 1♂ diss., Alanya, 10 km west [36.6°N, 31.89°E], 18-VII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T18, TAUI [2003-04]; 1♀ diss., Anamur, 30 km west [36.0845°N, 32.6082°E], 17-VIII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T17, TAUI [2003-09]; 1♀ diss., Ankara [39.927222°N, 32.864444°E], V-1937, Dr. Vasvari, D. elongata det. K. Lopatin 1961, HNHM [2004-08]; 1♂ diss., 2♀♀, Bergama, 5 km north [39.1553°N, 27.1629°E], 24-VIII- 1972, [on T. smyrnensis (Gerling and Kugler 1973)], T38, TAUI [2003-06]; 3♂♂ diss., 1♀ diss., 1♂, 1♀, Bürücek [Burucek; 37.35°N, 34.83333°E], Toros [Mt.], Anat[olia], 29–31-VII-1947, Exp. N. Mus. CSR,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 49 NMPC [2004-08, 2005-07, 14, 15]; 1♀ diss., 1♂, Cordélio [Karsiyaka; 38.4614°N, 27.1119°E], D. elongata det. V. Laboissière 1939, IRSNB [2006-10]; 1♂ diss., 2♀♀, Ephesos [Selcuk; 37.9517°N, 27.3747°E], 18-V- 1992, V. Nemec, EGRC [2005-03]; 1♂ diss., 2♂♂, Ereckli [Erekli; 41.28944°N, 31.41805E], Asia Minor, Bodemeyer, 824 [ZMHB], D. elongata [ZMHB], ZMHB [2003-08], NMPC; 1♂ diss., 2♀♀ diss., Eski–Chehir [Eskisehir; 39.77667°N, 30.52056°E], Asia Minor, Bodemeyer, NMPC [2004-29; 2005-29, 30]; 1♀ diss., Izmir, 10 km east [38.43°N, 27.19°E], 23-VIII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T41, TAUI [2003-08]; 1♂ diss., Kash [Kas], 30 km west [36.27°N, 29.333333°E], 20-VIII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T27, TAUI [2003-01]; 1♂ diss., Kassaba [Kasaba; 36.31111°N, 29.73472°E], Manisa Prov., Asia Minor, 1-VIII-1931, B.P. Uvarov, Brit. Mus. 1931-468, USNM [2004-01]; 1♂ diss., Malatya [38.353333°N, 38.311944°E], VI-1964, Seidenstücker, UH.G, DEI [2003-16]; 1♂ diss., 2♂♂, Ortakche [Ortakoy; 37.4167°N, 28.7167°E], east of Aydin on Menderes River, Asia Minor, 23-VII- 1931, B.P. Uvarov, British Museum 1931-468 [ZIN], BMNH [2003-09], ZIN [1♂]; 2♀♀ diss., Selcuk, 10 km east [37.8871°N, 27.4106°E], 22-VIII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T37, TAUI [2003- 05, 07]; 2♂♂ diss., Smyrna [Izmir; 38.407222°N, 27.150278°E], 1955, Erwerb, Coll. Brancsik, NHMB [2003-09, 11]; 1♂ diss., 5♀♀, Tarsus [36.91778°N, 34.89167°E], Asia Minor, Sterba, NMPC [2004-47]; 1♂ diss., 1♀ diss., 1♀, Turgutli [Turgutlu; 38.5008°N, 27.7058°E], 65 km east Izmir, 23-VIII-1972, [on T. smyrnensis (Gerling and Kugler 1973)], T40, TAUI [2003-02, 2004-01]; 1♂ diss., 1♀, Ushak, Abide [Usak; 38.68°N, 29.40805°E], 3-VI-1989, Kuff & Szailles, RBCN [2003-01]; 1♀ diss., Yeniköy [multiple possible geocoordinates], Toros [Mt.], Anat[olia], 30-VIII-1947, Exp. N. Mus. CSR, NMPC [2005-06]; UKNOWN COUNTRY: 1♀ diss., Dobrudscha [Dobruja region; around border of Bulgaria and Romania by Black Sea; specific locality not given; app. location: 43.6843°N, 28.5364°E], D. elongata, 126, Coll. Schultheiss, DEI [2005-06]; 1♀ diss., Tichakir [geocoordinates not locatable], Tal Noda, Asia Minor [possibly an archaeological site near the coast of Turkey], J. Weise collection, ZMHB [2003-07]; UNITED STATES OF AMERICA (introduced): California: Yolo Co.: 1♂ diss., Cache Creek near Rumsey [38.8895°N, -122.2333°W], 28-IX-2006, J. Herr, on T. parviflora, voucher [source: 3 km west Sfakaki (nr Stavramenos), Greece], GSWRL [2007-47]; Texas: Howard Co.: 1♂ diss., 4♂♂, 7♀♀, Higgins Ranch near Beals Creek, 32.251[069]°N, -101.386[506]°W, 8 km east of Big Spring, 18-VIII-2005, J.L. Tracy, on T. chinensis/T. canariensis [on regrowth following July defoliation by Diorhabda], voucher [source: 3 km west Sfakaki (nr Stavramenos), Greece], GSWRL [2006-11]; 5♂♂ diss., Higgins Ranch near Beals Creek, 32.24966°N, -101.38582°W, Big Spring, 1-IX-2006, J.L. Tracy, on Tamarix chinensis/T. canariensis, GSWRL [2008-01, 02, 03, 04, 05]; Potter Co.: 1♂ diss., 9♂♂, 5♀♀, Lake Meredith National Recreation Area, 35.52774°N, -101.76322°W, 11-IV-2006, V. Carney, on T. ramosissima, voucher [source: Posidi Beach, Greece], GSWRL [2006-18]. Distribution. General. Diorhabda elongata was most frequently collected from Italy to Bulgaria and central Turkey, but it occurs sporadically elsewhere around the Mediterranean in Lebanon, Egypt, Algeria, Portugal and Spain, and near the Caspian Sea in southern Russia (Dagestan Republic). Its native distribution is restricted to countries bordering on the Mediterranean Sea and the additional countries of Portugal, Bulgaria, Macedonia, and Russia (Map 2). Previous reports for the general distribution of D. elongata (Weise 1924, Heyden et al. 1891, Winkler 1924–1932, Ogloblin 1936, Wilcox 1971, Warchalowski 2003, Lopatin et al. 2004) are accurate only in their inclusion of the Mediterranean region to southern Russia. Further collections should provide specific localities of D. elongata along coastal areas of Portugal, France, Romania (Dobruja region), Syria, Algeria, and southern Russia (Dagestan Republic). Additional collections could also reveal D. elongata as common in Slovenia and uncommon in , Georgia and the coastal areas of Azerbaijan, Tunisia, Libya and Morocco. The D. elongata group is not reported from Israel (Lopatin et al. 2003). Confirmed Records. We have dissected specimens of D. elongata from the following countries with previous literature records (Map 2): Portugal (Corréa de Barros 1924), Spain (Hopkins and Carruth 1954, Torres Sala 1962, Petitpierre 1988), Italy (Reiche and Saulcy 1858, Weise 1893, Porta 1934, Zocchi 1971, Lundberg et al. 1987b, Biondi et al. 1995), Greece (Brullé 1832, Reich and Saulcy 1858, Regalin 1997),

50 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Bulgaria (Tomov 1969, 1979, 1984; Warchalowski 1974; Gruev and Tomov 1986, 1998), Turkey (Reiche and Saulcy 1858; Ogloblin 1936; Kerville 1939; Gerling and Kugler 1973 [list on p. 20 should indicate Turkey (T), not Israel (blank) beside D. elongata]; Aslan et al. 2000; Gök and Çilbiroğlu 2003, 2005; Gök and Duran 2004), Russia (Ogloblin 1936), Cyprus (Georghiou 1977), Syria (Reiche and Saulcy 1858, Ogloblin 1936), Lebanon (Reiche and Saulcy 1858 [as Syria], Ogloblin 1936 [as Palestine], Lopatin 1967 [as Lebanon]), Egypt (Reiche and Saulcy 1858, Weise 1925), Algeria (Reiche and Saulcy 1858, Peyerimhoff 1926), and the United States (Texas, California; introduced; Map 7, see Potenial in Tamarisk Biological Control below for more details) (Riley et al. 2003, DeLoach et al. in prep., Hudgeons et al. 2007a). We dissected D. elongata from a series of Beirut, Lebanon, identified as D. elongata by Lopatin (1967). A male D. elongata we dissected was collected in 1982 by Thure Palm from Fiumi di Pollina, Sicily, Italy, and it probably is associated with studies Palm co-authored reporting collections by himself and others (Lundberg et al. 1987a) of D. elongata on T. gallica at Fiumi di Pollina (Lundberg et al.1987b). New Records. We have dissected D. elongata from the following countries for which we find no previous specific reports in the literature: Croatia, , Montenegro, Macedonia (The Former Yugoslav Republic), and Albania. Unconfirmed Records. We cannot confirm reports of D. elongata from Georgia (Reiche and Saulcy 1858, Lozovoi 1961) and Azerbaijan (Samedov and Mirzoeva 1985; Mirzoeva 1988, 2001). Because of the predominance of D. carinata in these areas, we consider these reports to primarily involve D. carinata (see D. carinata—Distribution below), and the presence of D. elongata needs further confirmation. However, we have dissected D. elongata from the Dagestan region of Russia along the Caspian Sea (Fig. 19—Dagestan) and suspect that it is present in Georgia and Azerbaijan. Reports of D. elongata in areas east of the Caspian Sea in central Asia (east of 50°E) (e.g., Lopatin, 1977a, Medvedev and Voronova 1977b, Bieńkowski 2004) should refer instead to D. carinata and D. carinulata. France lies between two countries with confirmed host records, Italy and Spain (Map 2). Therefore, although we have not examined D. elongata from France, we still consider the general locality record of France as accurate (Reiche and Saulcy 1858, Bedel 1892, Laboissière 1934, Chatenet 2002). All locations from which specimens were dissected in Bulgaria (9), Greece (49; with the exception of a single male specimen of D. carinata considered as mislabeled from Attica, Greece), and Italy (15) were D. elongata and we consider all literature records of D. elongata from these countries as accurate (Map 2). We examined a specimen of D. elongata from Italy (ZMUH) with an identification label of D. e. ab. carinata by Laboissière and we consider Porta’s (1934) report of D. e. ab. carinata in Italy to be D. elongata. In Turkey, D. carinata is found as far west as 41.5°E and it is found near the south Turkish border in Halab, Syria at 37°E longitude (Map 3). Therefore, we are only confident in accepting literature records of D. elongata in Turkey that are in the area west of 35°E longitude, from which we dissected D. elongata from all 21 available locations (Map 2). Below are 13 unconfirmed locality records of D. elongata that we consider as valid (Map 2): BULGARIA: environs, near Struma River [42.01670°N, 23.08000°E], on Tamarix (Tomov 1979); 1 specimen, Melnik, 4-V-1971, Stück (Warchalowski 1974) [D. elongata dissected from same location]; GREECE: Kaiafas [37.5167°N, 21.6000°E], Peloponnes [Pelopónnisos Peninsula], west coast, 24- V-1995 (Serge Doguet, Fontenay–sous–Bois, France, pers. comm.); Plakias [35.200°N, 24.400°E], Crete, 13- 16-VI-1995; Rethimnon nom. [Rethymnon; 35.3647°N, 24.4714°E], Crete, 13-16-VI-1995; Sèlia [Sellia; 35.4000°N, 24.2330°E], Crete, 13-16-VI-1995, (Regalin 1997; on Tamarix smyrnensis); ITALY: Fiume de Pollina [38.01667°N, 14.6667°E], 15 km east Cefalu, Sicily, on Tamarix gallica [D. elongata dissected from same location]; Fiume de Tusa [38.01667°N, 14.2667°E], 21 km east Cefalu, Sicily, on Tamarix gallica (Lundberg et al. 1987b); Trieste environs [45.6395°N, 13.7876°E] (Weise 1893); TURKEY: Asagigökdere village [37.59722°N, 30.82833°E], Isparta Province, 650 m [elev.], 12-V-2000 (2♂♂, 3♀♀), on Tamarix smyrnensis, 2-VII-2000 (3♂♂, 4♀♀), 10-X-2000 (1♂, 1♀), 4-V-2001 (3♂♂, 3♀♀) (Gök and Çilbiroğlu 2003, Gök and Duran 2004); 4 specimens, Edirne [41.67440°N, 26.56080°E], 15 m [elev.], 6-V-1960 (Tomov and Gruev 1975); 10 specimens, Yozgat dintorm [39.8200°N, 34.8044°E], Yozgat [Ili], 1,300 m [elev.], 26- VI-1975, G. Osella (Tomov 1984).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 51 Because of the proximity to locations of D. carinata, the following two unconfirmed distribution records of D. elongata from east of 35°E in Turkey are uncertain (Map 2): TURKEY: 1 specimen, Diyarbakir dintorm [37.9189°N, 40.2106°E], Diyarbakir [Ili], 5-VI-1971, G. Osella; 5 specimens, Bafra [41.56780°N, 35.9069°E], Samsum [Ili], 8-VII-1975, G. Osella (Tomov 1984). Discussion. Taxonomy. Diorhabda elongata (Brullé) was described in the genus Galeruca by Brullé (1832) from Morée (= Pelopónnisos peninsula), Greece. We dissected a total of 16 male and 11 female topotypes from 12 locations of the type locality, Pelopónnisos, Greece (Map 2). These males and females shared unique genitalic characters among the D. elongata group, comprising a distinct genitalic morphotype pair along with all other specimens examined from Greece and the surrounding countries of Italy, Albania, Macedonia and Bulgaria as well as western Turkey (see discussion below; Figs. 14, 19, 24, 29, 34, 39, 44; Map 2). We are certain that this genitalic morphotype pair is a single species conspecific with D. elongata. The endophallic sclerites of D. elongata bear several characters distinguishing them from those of the holotypes of D. carinata, D. carinulata,andD. meridionalis (as illustrated in Figs. 18–19 of Berti and Rapilly 1973) and other specimens of these species and D. sublineata examined in this revision (see Male-Genitalia above; Figs., 15–18, 20–23, 25–28, 30–33). Additional unique qualitative characters of the female genitalia distinguish D. elongata from other species in the D. elongata group (see Female-Genitalia above; Figs. 34–43). The distinctive genitalic characters of D. elongata are maintained in the same areas where D. sublineata, D. carinata and D. carinulata occur and near its abutting range boundary with D. meridionalis, and this is strong evidence for reproductive isolation between these species (see Biogeography below; Map 1, Table 8). Further evidence for reproductive isolation between D. elongata and several members of the D. elongata group is also found in previously discussed differences in component ratios of putative aggregation pheromones and reduced F2 hybrid egg viabilities. Both the text and color habitus drawing accompanying Brullé’s (1832, Plate 44, Fig. 10) original description lack any indication of striping or vittae on the elytra, but we find that D. elongata commonly has submarginal and substural vittae (Fig. 1). The elytral vittae in D. elongata are confined to the apical half of the elytra (Fig. 1), distinguishing them from some specimens of the other four species of the D. elongata group in which the elytral vittae, when present, may extend into the basal half of the elytra (Figs. 5, 9). We find that external characters previously used to distinguish D. elongata from the sibling species D. carinata and D. sublineata (Weise 1883, 1890; Laboissière 1934, Porta 1934, Bechyné 1961, Warchalowski 2003) are too variable for species diagnosis (for further details, see Discussion—Taxonomy under D. carinata and D. sublineata). We have dissected specimens of D. elongata that were misidentified by taxonomists using external diagnostic characters as D. e. ab. carinata in Italy, D. e. var. sublineata in Greece, and D. e. ab. sublineata in Italy (see Material examined). Gruev and Tomov (1986) provide a detailed description of external adult morphological characters for D. elongata but their given size range of 4.5–8 mm, is wider than the size range we find of 5.3–7.7 mm, and their range certainly includes that of sibling species that are smaller (D. carinulata) and larger (D. carinata) (Table 2). The size range of 5.5–8 mm given elsewhere in southern Europe (Porta 1934, Laboissière 1934) is closer to our observation regarding minimum size. Mulsant (Mulsant and Wachanru 1852) described G. costalis Mulsant from Cilicia (along the coast of southwestern Turkey, towards Syria) from specimen(s) of unknown sex. The given body length is 5.6 mm, possibly making the type a male (see Table 2). The elytral vittae (as striae) are described beginning at a point 3/5th the length of the elytra from the base (starting in the apical half) and ending at a point 6/7th the length of the elytra from the base (ending near the apical tip). Confinement of the vittae to the apical half of the elytra agrees with our observations for D. elongata. We dissected D. elongata from all four locations available from the type locality of Cilicia (an area including Adana in southwest Turkey, Map 2). We dissected D. carinata and D. meridionalis from ca. 100 km southeast of Cilicia in Halab, Syria (Map 1). We follow Reiche and Saulcy (1858) in regarding G. c o st a li s as a synonym of D. elongata. Other synonyms established by Reiche and Saulcy (1858) and Weise (1893) actually consist of the three valid species D. carinata, D. sublineata, and D. carinulata.

52 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS The number of spines on the endophallic sclerites and the shape of the elongate sclerite are fairly variable in D. elongata (Fig. 19) and males from certain series were often more similar to one another than those from other series. For example, a majority of males dissected from Promontorio de Gargano, Italy, and 60 km E. Thessaloniki, Greece had greater spination of both the elongate and palmate sclerites than males from Burucek, Turkey. However, the variability in spination is continuous (see Figs. 19 and 29, Methoni, Greece), and we detected no geographic pattern. We have seen only a few series of specimens collected from identified species of Tamarix hosts and further investigations should be made into potential patterns in morphological variability associated with host species. Lohse (1989) found sympatric and phenologically differing Rumex and Polygonum host ecotypes of Galerucella aquatica that could be distinguished by the coloration of the last abdominal sternite but not by endophallic sclerites. Studies of genetic relationships among D. elongata populations collected from various Tamarix spp. in southern Europe are in progress (R. Carruthers, USDA/ ARS, Albany, CA, pers. comm.). Common Name. The name “Mediterranean tamarisk beetle” refers to the great majority (ca. 80%; Fig. 52B) of collections of D. elongata being from the Mediterranean biome. Biology. Host Plants. Diorhabda elongata has been collected from Tamarix smyrnensis Bunge, a close relative of T. ramosissima (Baum 1978), on Crete, Greece (Regalin 1997) and in Isparta Province of southwest Turkey (Gök and Çilbiroğlu 2003, 2005; Gök and Duran 2004) (Table 1). Table 5 of Gerling and Kugler (1973) lists T. smyrnensis as the host species for seven locations in western Turkey associated with dissected TAUI specimens of D. elongata (see above Material Examined). Tamarix gallica is reported as a host in northern Sicily (Lundberg et al. 1987b), and we dissected a specimen from the Toscana province of Italy with a host label of T. gallica. Diorhabda elongata is reported from Tamarix sp. in Cyprus (Georghiou 1977). New host records from collections on Peloponnisos, Greece by Javid Kashefi (USDA-ARS) are T. hampeana Boissier and Heldreich and T. parviflora. Tamarix parviflora is also a reported host from Cache Creek, California (DeLoach et al. in prep.; R. Carruthers, pers. comm.). Tamarix chinensis × T. canariensis/T. gallica is a new host record for populations of D. elongata established at Big Spring, Texas (Table 1). Dalin et al. (in press) found that D. elongata from Crete preferred T. parviflora to a similar degree as T. ramosissima in multiple-choice field cage studies in California. In Bulgaria, D. elongata severely defoliates Tamarix sp. trees (especially those in open areas) when it becomes numerous on the dry sandy terraces of the Struma River valley near Blagoevgrad (Tomov 1979). Pallas, a close relative of T. parviflora (Zieliński 1994), is common along the Struma River Valley and other parts of Bulgaria, Greece, and Turkey (Baum 1978, Zieliński 1994), where it may serve as a host for D. elongata. Tamarix dalmatica Baum, the prevalent Tamarix species along the eastern coastlands of the Aegean Sea (Baum 1978), may serve as a host in Croatia, Montenegro and Albania. Reports of D. elongata in Central Asia from the leguminous shrubs Halimodendron (Sinadsky 1960, Bieńkowski 2004), Ammodendron (Seitova 1974, Sinadsky 1968), and Alhagi (Sinadsky 1968) (Fabaceae) should refer to Galerupipla sp. (see DeLoach et al. 2003b). No-choice larval host suitability studies by Milbrath and DeLoach (2006a) confirm that D. elongata larvae from Crete can survive to adulthood only on plants of the order Tamaricales, including Tamarix (Tamaricaceae) and, to a lesser degree, on three North American Frankenia spp. (Frankeniaceae): F. s al in a , F. johnstonii, and F. ja m e s ii. Herr et al. (2006, in prep.) found larval survival on F. salina was not different than that on T. ramosissima. Multiple-choice adult oviposition studies in field cages revealed that the three North American Frankenia spp. provide little attraction for oviposition compared to Tamarix (Milbrath and DeLoach 2006a; Herr et al. 2006). Diorhabda elongata tended to oviposit less on T. aphylla compared to other invasive North American tamarisk, including T. ramosissima, T. chinensis, and T. canariensis/T. gallica in multiple choice field cage tests (Milbrath and DeLoach 2006a, Herr et al. 2006). However, in other multiple-choice field cage tests among Tamarix spp., oviposition by D. elongata on one accession of T. aphylla (Phoenix, Arizona) was not different than that on T. ramosissima × T. canariensis/gallica, T. canariensis/T. gallica and T. parviflora (Milbrath and DeLoach 2006b). In a no-choice laboratory cage experiment, D. elongata accepted T. aphylla for oviposition to a significantly lesser extent than T.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 53 ramosissima but not T. parviflora (Herr et al. 2006). In this same no-choice experiment, the difference between oviposition by D. elongata on F. salina (inland variety) and T. ramosissima, T. parviflora,and T. aphylla was not significant (Herr et al. 2006). Diorhabda elongata accepted T. aphylla for oviposition to the same degree they accepted T. ramosissima × T. chinensis in no-choice field cage studies (Milbrath and DeLoach 2006b). In open field testing with transplanted potted plants at Big Spring, Texas, D. elongata oviposited little on T. aphylla compared to T. ramosissima/T. chinensis (Moran et al. in press), and almost no oviposition occurred on F. salina (Herr et al. 2006). Tamarix aphylla is at moderate risk of damage by D. elongata in the field and it is difficult to predict to what degree D. elongata would damage T. aphylla, especially in the absence of other Tamarix spp. (Milbrath and DeLoach 2006b). Based on open-field testing in south Texas, Moran et al. (in press) concluded that D. elongata will likely have limited establishment and impact on T. aphylla. Frankenia is at very low risk of damage from D. elongata (Milbrath and DeLoach 2006a). Risk of damage to both T. aphylla and Frankenia by D. elongata is probably much lower when these plants are not in the proximity of preferred Tamarix spp. (e.g., Blossey et al. 2001). Ecology and Phenology. Tomov (1979) studied large populations of D. elongata from 1967–1971 along the Struma River near Blagoevgrad, Bulgaria. They report that the first eggs appear in early April and mating pairs are found from the middle of April until September. Larvae appear in early May until September, and the last adults are seen in October. Three population peaks of adults are seen near Blagoevgrad: late May and early June, late July and early August, and late September. Feeding by young larvae produces small holes in the lower epidermis and parenchyma of Tamarix leaves. Older larvae and adults eat entire leaves. Larvae pupate in loose cocoons in the soil. Late in the summer, many Tamarix shrubs, especially isolated ones, are completely defoliated along the Struma River by D. elongata. Diorhabda elongata was noted in small numbers from May to October on T. smyrnensis in southwest Turkey (Gök and Çilbiroğlu 2003). Adult collection dates from our examined material are from May to September in Croatia, April to October in Greece, and April to October in Italy. Our only specimens from Spain were collected on 15 February at Estepona on the southern coast. Milbrath et al. (2007) found that D. elongata from Crete overwintering at Temple, Texas had ca. 80–95% survival from early November through the middle of March when tamarisk leaves began budding. Overwintered adults emerged in mid to late March and commenced ovipositing at the beginning of April giving rise to four generations and a partial fifth generation. Fourth generation adults emerging in early September oviposited little in September and ceased oviposition by October when they appeared to enter diapause. In Big Spring, TX, five generations were also observed with overwintering adults generally emerging from late March and early April and fifth generation adults appearing from October to early November (DeLoach 2008, DeLoach et al. in prep.). On the Rio Grande from Candelaria to near Presidio, TX, large numbers of adults were actively defoliating trees into mid-November (A. Berezin, Sul Ross State University, Alpine, TX, pers. comm.). From Big Spring during 2005–2008, data was collected on the population density of D. elongata and associated rate of tamarisk defoliation on trees along sample transects throughout the growing season to develop models of wave dispersal using both a mathematical deterministic model and a statistical spatial regression model (J. Sanabria, pers. comm., DeLoach 2008, and DeLoach et al. in prep.). Development and Reproduction. Egg mass size in the field in Bulgaria ranged from 10 to 19 eggs (26 in the laboratory) (Tomov 1979). In laboratory studies at 28°C by Milbrath and DeLoach (2006b), D. elongata produced 16.0 ± 0.5 eggs per mass and fecundity averaged 219 ± 56.2 eggs with a population doubling time of 5.8 days on T. ramosissima × T. chinensis. On T. aphylla, fecundity was higher at 328.0 ± 60.4 but the population doubling time was longer at 6.6 days, which was partly due to both longer preoviposition and oviposition periods on T. aphylla. Milbrath et al. (2007) compared D. elongata from Crete with D. carinulata, D. carinata and D. sublineata at 28°C, and found D. elongata was similar to the other species with a development time of 21.0 days from egg to adult (with 78% survival), a fecundity of 281 eggs, and a population doubling time of 6.2 days.

54 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Natural Enemies. Tomov (1974) reported the tachinid Erynniopsis antennata (Rondani) (= E. rondanii Townsend) emerging from third instar larvae of D. elongata collected on 10 July in Bulgaria. E. antennata is also reported from southern France and Italy, and it also attacks the , Xanthogaleruca luteola (Müller) (Tschorsnig and Herting 1994). Adult E. antennata oviposit on beetle larvae and tachinid adults emerge from late stage third instar larvae (prepupae) during the growing season and from overwintering beetles in the early spring (Dreistadt and Dahlsten 1990). In 1939, E. antennata was introduced from Europe and established in California for control of the exotic elm leaf beetle (Flanders 1940). The parasitic has now spread throughout California (Dahlsten et al. 1998), and is also reported from Oregon (USNM collection data). In California, parasitism rates of overwintering elm leaf beetle adults can reach 65%, but the fly is limited by a eulophid hyperparasite Baryscapus erynniae (Domenichini) (Dreistadt and Dahlsten 1990). A protozoan mircorsporidian, Nosema sp., was found in adult D. elongata originating from near Sfakaki, Crete, Greece (shipment EIWRU-2002-1002) (identified by J. Siegel) and Posidi Beach, Greece (shipment EIWRU- 2002-1009) (D. Bean, pers. comm.). Biogeography. Comparative. Diorhabda elongata differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) strongly maritime and generally found within ca. 250 km from a sea coast, under 1,400 m elevation; (2) usually found in warm temperate Mediterranean woodlands or Temperate Broadleaf and Mixed forest biomes; and (3) latitudinal range of 30–45°N and most common from 35–43°N (Table 7, Figs. 51–52). Diorhabda sublineata is marginally sympatric with D. elongata in Portugal, Spain and Egypt and both occur in the Mediterranean Forests, Woodlands and Scrub biome around the Mediterranean Sea (Map 1, Tables 8 and 9, Fig. 52B). However, D. elongata predominates in the northeastern Mediterranean in Europe and Asia while D. sublineata predominates in the western Mediterranean and all of North Africa, including more xeric biomes such as desert and flooded grasslands in which D. elongata is rare or absent. Diorhabda sublineata additionally differs from D. elongata in being common further south at 31–35°N and ranging much further south to 16°N (Figs. 51–52). Diorhabda are unreported along two ca. 600–700 km stretches of land around the Mediterranean where the main distributions of D. elongata and D. sublineata should interface in western Italy and Israel and Palestine, and no D. sublineata are reported from the main area of distribution of D. elongata (Map 1). If these D. elongata/D. sublineata interface zones actually lack Diorhabda populations, it may be worth investigating the potential existence of some form of localized competitive mutual exclusion between these species. Diorhabda elongata is probably marginally sympatric with D. carinata in eastern Turkey, western Syria, and, possibly, southern Russia, Georgia and Azerbaijan. Diorhabda elongata is marginally sympatric with D. carinulata in southern Russia (Dagestan) (Map 1; Table 8). Both D. carinata and D. carinulata are strongly continental and are mostly found in desert and grassland biomes from which D. elongata is not known. Diorhabda meridionalis is apparently parapatric with D. elongata in Syria and differs from D. elongata in being common both in deserts and further south at 26–31°N (Tables 7 and 8; Map 1; Figs. 51–52). Descriptive. Diorhabda elongata is found in the west-central Palearctic realm (Maps 1). It is primarily collected from 35–43°N in the northeastern Mediterranean from Italy to Bulgaria and western Turkey in two biomes: the Mediterranean Forests, Woodlands and Scrub and the Temperate Broadleaf and Mixed Forests (Map 2). Ecoregions (Olson and Dinerstein 2002) inhabited by D. elongata from 35–43°N in the Mediterranean Forests, Woodlands and Scrub biome, based on frequency of collection, are the Ilyrian Deciduous Forests from Croatia to western Greece, the Crete Mediterranean Forests, and the Aegean and Western Turkey Sclerophyllous and Mixed Forests from coastal Greece to Turkey (Map 2). The latter is the center of distribution for T. hampeana (Boratyński et al. 1992, Zieliński 1994), a host plant of D. elongata. Tamarix hampeana occurs only rarely in the western Mediterranean area (De Martis et al. 1986), similar to D. elongata (Map 2). The primary range of another host of D. elongata, T. parviflora, is also in the eastern Mediterranean (Map 2). In the Temperate Broadleaf and Mixed Forests biome, D. elongata inhabits the Balkan Mixed Forests from Macedonia to northwest Turkey. This area is a possible source for a putative northern interior lowland D. elongata climatype occurring from 41–42°N and 200–500 m elev.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 55 Potential in Tamarisk Biological Control. Summary. The Mediterranean tamarisk beetle is providing effective biological control of T. parviflora at Cache Creek, California and Tamarix ramosissima/T. chinensis near Big Spring and Pecos, Texas (Map 7). Based on its biogeographic characteristics, D. elongata is most suitable for Mediterranean biome of northern California (Figs. 51–52; Map 13). D. elongata from Crete readily accepts the novel host T. chinensis × T. canariensis/T. gallica at Big Spring, Texas, and its absence in the deserts and grasslands of central Asia is probably due to poorer adaptation to the bioclimatic conditions of deserts and grasslands compared to other Diorhabda species rather than the lack of suitable Tamarix hosts. Diorhabda elongata will establish in west Texas grasslands and deserts around 31–32°N, but our HSI model predicts other Diorhabda species are better biogeographically suited to deserts and grasslands of the southwestern U.S., including Diorhabda carinulata, D. carinata, and D. sublineata. More adapted Diorhabda species may eventually replace D. elongata where it establishes in desert and grassland habitats (Map 13). Discussion. The Mediterranean tamarisk beetle attacks and defoliates Tamarix (Tomov 1979) in Temperate Broadleaf and Mixed Forest biome at ca. 42°N in Bulgaria. Tamarix smyrnensis is a close relative of the invasive T. ramosissima in North America (Baum 1978, Zieliński 1994), and it is attacked by D. elongata in Greece and Turkey. Diorhabda elongata attacks and damages both invasive T. parviflora and T. ramosissima/T. chinensis in North America. It has a moderate risk of damaging T. aphylla (Milbrath and DeLoach 2006b) and very low risk of damaging Frankenia (Milbrath and DeLoach 2006a), and both these risks are probably much reduced at less proximity to preferred Tamarix spp. (e.g., Blossey et al. 2001). The Mediterranean tamarisk beetle may be best adapted for areas from 35–43°N in the maritime temperate warm Mediterranean Forests, Woodlands and Scrub biome and the Temperate Broadleaf and Mixed Forests biome (see Biogeography). Of these two biomes, only the Mediterranean Forests, Woodlands and Scrub biome is found in the western U.S. From 35–43°N, the Mediterranean biome is represented by the California Interior and Woodlands and the California Montane Chaparral and Woodlands ecoregions (Map 13). The Mediterranean species T. parviflora is the dominant invasive tamarisk in these ecoregions (Map 7). Diorhabda elongata from Crete has established well on T. parviflora near Cache Creek, California (39°N, Map 7), where it entirely defoliated over 200 hectares of tamarisk along an ca. 40 km reach in 2007, and over ca. 250 ha. (600 acres) along 50 km (including parts of nearby Bear Creek) in 2008 (DeLoach et al. in prep.; R. Carruthers, pers. comm.). This estimate of defoliated area is based upon an analysis of aerial photography which revealed ca. 396 ha of T. parviflora along a similar 40 km reach (ca. 10 ha of tamarisk per creek km) of Cache Creek in 2001 (Ge et al. 2006). Tamarisk is estimated to occur at about half this density (ca. 5 ha. per km) over most of Cache and nearby Bear creeks where Diorhabda is well established (R. Carruthers, pers. comm.). Initial establishment at Cache Creek was slow, with very low populations persisting from 2004–2005, and only ca. 1.4 ha defoliated in 2006, and D. elongata has only weakly established and not established at some other northern California sites (John Herr, USDA/ARS, Albany, CA, pers. comm.). The possibility that establishment of D. elongata was slowed by predators or the need to adapt to a colder climate and to colder weather starting at longer daylengths earlier in the season should be investigated. At the Cache Creek site, potential parasitism of D. elongata by the previously discussed exotic parasitoid tachinid fly Erynniopsis antennata is being monitored (J. Herr, pers. comm.). Diorhabda elongata from Crete is also well established on T. chinensis × T. canariensis/T. gallica (identified by J. Gaskin) in the Temperate Grasslands, Savannas and Shrublands biome (Western Short Grasslands ecoregion) near Big Spring, Texas (32°N, Map 7) (Hudgeons et al. 2007a). During the first year of establishment at Big Spring in 2004, about four small trees were defoliated throughout the season, the first two trees being defoliated in July. In 2005, field populations increased and, by late September, over 200 tamarisk trees were totally defoliated, representing 0.17 ha of tamarisk canopy covered over a 0.66 ha area. By late October 2006, tamarisk over an ca. 7 ha area was defoliated (Everitt et al. [2007] classified defoliation from remote sensing by color aerial photography), ca. 13 ha was defoliated in 2007, and over 40 ha was defoliated in 2008, including defoliation of an ca. 7 km stretch of Beals Creek with satellite populations defoliating scattered trees over a 19 km wide area around Big Spring. At each of two sites in the Trans-Pecos Chihuahuan Desert on the Pecos River near Pecos and Imperial, Texas (Map 7), ca. 1 ha of tamarisk trees

56 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS were defoliated in 2007 during the second year post release (Mark Muegge, The Texas AgriLIFE Extension Service, Fort Stockton, TX, pers. comm.). By November 2008, defoliation by D. elongata along the Pecos River north of Pecos progressed to ca. 3.2 river km, but no defoliation was found near Imperial. Establishment was initially marginal or unsuccessful at other Texas AgriLIFE release sites in west Texas, but the use of ant baits in 2008 appears to be improving establishment at several of about 16 additional sites (A. Knutson, pers. comm.; see Map 7). Releases of populations of D. elongata from Crete were first made along the Rio Grande in Texas at several sites from Candelaria towards Presidio (Tyrus Fain, Rio Grande Institute, La Junta Project, Marathon, TX, pers. comm.) in the summer of 2007. Releases were made again in these areas in 2008 and in Big Bend National Park (Joe Sirotnak, Big Bend NP, TX, pers. comm.) and nearby Adams Ranch (M. Muegge, pers. comm.). Partial defoliation by D. elongata of several trees surrounding the release cages were noted at three sites between Presidio and Candelaria in the late summer and fall of 2008 (Mark Donet, USDA- NRCS, Alpine, TX, and Andrew Berezin, pers. comm.). In 2008, further releases were also made at various sites in west Texas, such as Iraan and Twin Buttes Reservoir near San Angelo (A. Knutson, pers. comm.), and at Holloman AFB, New Mexico (D. Thompson, pers. comm.). In the Western Short Grasslands at Lake Meredith in north Texas (35°N, Map 7), D. elongata from Posidi Beach, Greece (40°N), overwintered in the open field after its release in 2005 and produced spotty damage to tamarisk, dispersing about 1 km, but it was present in only small numbers in 2008 (Erin Jones, Texas AgriLIFE Research, Amarillo, TX, pers. comm.). D. elongata from Crete failed to establish at Seymour, Texas in the Central and Southern Mixed Grasslands in 2004, but D. carinata from Qarshi, Uzbekistan was released there in 2008 and appeared to be establishing, defoliating over 0.2 ha during the first year of release (C. Randal, pers. comm.). In 2003, populations of D. elongata from Crete were released in the Chihuahuan Desert near Artesia, New Mexico (33°N, Map 7) and initially increased in 2004, but severely declined in early 2005 and could not be found through 2008. Although the Mediterranean tamarisk beetle appears to be very promising for tamarisk biocontrol across the southern U.S. (Milbrath et al. 2007), it has never been recorded in its native habitat from two biomes: Deserts and Xeric Shrublands and Temperate Grasslands, Savannas and Shrublands (Table 9; Fig. 52B). We are not aware of a desert or grassland ecotype of D. elongata. During 2004–2007, D. elongata defoliated ca. 13.8 ha in the grassland biome at Big Spring compared to ca. 200 ha in the Mediterranean biome on Cache Creek. If the trend continues for more rapid defoliation in California, this could further support our biogeographic models showing that the Crete beetle is best suited to Mediterranean biomes. Other species of tamarisk beetles are common in desert and grassland biomes (Map 1, Table 9), and they may be better suited to much of the large area of these biomes invaded by tamarisk in North America as estimated by our relative Habitat Suitability Index Models (Map 13). Species distribution models incorporating climatic data are planned to better evaluate the potential suitability of D. elongata for climates across the western U.S. Making accurate predictions regarding potential North American ranges of any Diorhabda could be complicated if their field host preferences for Tamarix spp. vary significantly across their distributions.

Diorhabda carinata (Faldermann, 1837) larger tamarisk beetle (Figs. 3, 4, 15, 20, 25, 30, 35, 40)

Galeruca carinata Faldermann, 1837:329 (Type locality: Transcaucasus region [Georgia, Armenia and Azerbaijan]; as Galleruca). Galeruca elongata: Reiche and Saulcy, 1858:42 (part, l’Immerétie [western Georgia], Syria, Turkey, as Galleruca); Joannis, 1866:83 (part, monograph; Syria, as Galleruca). Diorhabda elongata: Weise, 1883:316 (part, established genus; Transcaucasus); Holdhaus, 1920:45 (Assur [Ash Shar- qat], Iraq); Ogloblin, 1936:79 (part, Iran, Syria, Turkey, Transcaucasus, central Asia); Rusanov, 1949:118 (part, Cen- tral Asia; as Diorrhabda); Kyrzhanovskiy, 1952:198 (part, Turkmenistan), 1965:392 (part, middle Asia); Pavlovskii and Shtakelberg, 1955:566 (part, southwest Europe, as Diorrhabda); Yakhontov and Davletshina, 1955:58 (part,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 57 biology; Amu Darya Delta, northern Uzbekistan); Sinadsky, 1957:950, 1963:84, 1968:64 (part, biology; Amu Darya Valley, Uzbekistan; as Diorrhabda); Medvedev, 1959:118 (part, Turkmenistan); Yakhontov, 1959:338 (part, biology; Uzbekistan); Lozovoi, 1961:86 (part, eastern Georgia); Kulinich, 1962:73 (part, biology; Tajikistan); Pripisnova, 1965:83 (part, biology, Tajikistan); Kulenova, 1968:171 (part, southeastern Kazakhstan); Lopatin and Tadzhibaev, 1972:591 (part, Tajikistan); Lopatin, 1977a:282 (part, Asia), 1981:375 (Robate–Ghozlog [Robat–e Qozlog], Iran); Davletshina et al., 1979:79 (part, southwest Kyzyl–Kum Desert, central Uzbekistan); Habib and Hasan, 1982:19 (host range; northern Pakistan); Tomov, 1984:377 (part, Artvin, Turkey); Samedov and Mirzoeva, 1985:712 (part, Azerbaijan); Mirzoeva, 1988:?, 2001:48 (part, Azerbaijan); Lopatin and Kulenova, 1986:129 (part, Kazakhstan); Myartseva, 1995:4, 1999:1; 2001:1 (part, biology; Turkmenistan); Kovalev, 1995:78 (part, south-central Palearctic); Richter and Myartseva, 1996:316 (parasitoid, Turkmenistan); Gruev and Tomov, 1998:70 (part, Transcaucasus, mid- Asia); Aslan, 1998:287 (Ezurum Ili, province of northeast Turkey); Aslan et al., 2000:30 (part, eastern Turkey); Anonymous, 2001:52N (part, Turkmenistan); DeLoach et al., 2003a:230 (part, Uzbekistan [Qarshi]), 2003b:126 (part, host range; Pakistan, Uzbekistan, Turkmenistan, Georgia, Azerbaijan), (2008, in prep.) (part, Qarshi, Uzbeki- stan); Khamraev, 2003:11 (part, Uzbekistan); Milbrath et al., 2003:225 (part, Qarshi, Uzbekistan); Riley et al., 2003:69,189 (part, catalog of North America [introduced]); Warchalowski, 2003:328 (part, taxonomic keys; Cauca- sus and Central Asia); Bieńkowski, 2004:76 (part, keys; eastern Europe, Caucasus, Iran, Iraq, Central Asia, Kazakh- stan); DeLoach and Carruthers, 2004a:13, 2004b:311 (part, Uzbekistan [Qarshi]); Gök and Duran, 2004:17 (part, Turkey); Lopatin et al., 2004:127 (part, central Asia); Gates et al. 2005:28 (parasitoid; Ashgabat, Turkmenistan); Dudley, 2005a:13, 2005b:42N (part, biological control; ex: Uzbekistan); Milbrath and DeLoach, 2006a:32, 2006b:1379 (part, host specificity; Qarshi, Uzbekistan); Dudley et al., 2006:137 (part, host range; Qarshi, Uzbeki- stan); Milbrath et al. (2007) (part, biology; Qarshi, Uzbekistan); DeLoach (2008); Bean and Keller (in prep.) (part, diapause induction; Qarshi, Uzbekistan); Thompson et al. (in prep.) (part, laboratory hybridization; Qarshi, Uzbeki- stan). Diorhabda elongata var. carinata: Heyden et al., 1891:375 (part, catalog for Europe and Caucasus); Weise, 1893:635, 1132 (part, Transcaucasus); Jacobson, 1901:137 (part, Amu Darya (river), Turkmenistan, as Dirrhabda). Diorhabda elongata ab. carinata: Weise, 1924:78 (part; world catalog; Transcaucasus; Turkmenistan, Amu Darya); Win- kler, 1924–1932:1307 (part, Palearctic catalog, Transcaspian region); Warchalowski, 2003:328 (part, taxonomic keys, Caucasus and Central Asia). Diorhabda rybakowi: Mityaev, 1958:86 (part, biology, Kazakhstan, as rybakovi). Diorhabda elongata carinata: Bechyné, 1961:256 (Pol–e khomri [Polichromi], Afghanistan; central Asia); Lopatin, 1963:355 (Afghanistan); Wilcox, 1971:63 (world catalog, Afghanistan); Medvedev, 1983:123 (, Afghanistan); 1985:44 (Afghanistan); Lopatin et al., 2004:127 (east Kazakhstan, northwest China); Dalin et al. (in press) (host range; Qarshi, Uzbekistan). Diorhabda elongata sublineata: Gressitt and Kimoto, 1963a:407 (part, Asia Minor, W. Asia); Wilcox, 1971:63 (part, world catalog, Asia Minor, W. Asia). Diorhabda carinata: Berti and Rapilly, 1973:881 (restored species, Transcaucasus), DeLoach et al., 2003b:126.

Male. Genitalia. Diorhabda carinata can be distinguished from all other members of the D. elongata group by a combination of characters of the palmate endophallic sclerite (PES) and the lack of a connecting endophallic sclerite (CES). The palmate endophallic sclerite (PES) in D. carinata always bears a strong lateral appendage (LA) and the distal margin is truncate-serrate with two to six (commonly four to five) usually distal spines, a maximum of one spine being subdistal (Figs. 15, 30). In contrast, the PES of D. carinulata (Figs. 17, 32) and D. meridionalis (Figs. 18, 33) lacks a lateral appendage (but may bear a lateral notch) and the distal margins of the PES are narrowly rounded and generally smooth with one or two small subdistal spines that sometimes project beyond the distal margin. The PES of D. elongata also lacks a lateral appendage and is usually rounded with mostly subdistal spines and a maximum of two distal spines (Figs. 14, 29). Diorhabda sublineata bears a CES connecting the PES to the elongate endophallic sclerite (EES) (Figs. 16, 21, 31), but the CES is lacking in D. carinata (Figs. 15, 20, 30) (in some darkly sclerotized specimens of D. carinata, a faint lateral line is seen where a CES would be found). In D. carinata, the spined area of the EES extends greater than or equal to 0.34 times the length of the EES (Figs. 15, 20). In contrast, the spined area of the EES is confined to less than or equal to 0.16 times the length of the sclerite in D. elongata (Table 3; Figs. 14, 19, 24, 48). In D. carinata, spines of the EES are often irregularly spaced along the blade with conspicuous gaps (Fig. 20). In D. carinulata and D. meridionalis, spines along the EES are usually evenly and closely spaced along the blade (Figs. 22–23). The EES of D. meridionalis additionally bears a hooked apex that is absent in D. carinata.

58 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Measurements. See Tables 2 and 3. Female. Genitalia. Female D. carinata may be distinguished from all other members of the D. elongata group except D. sublineata by their triangulate vaginal palpi (VP) that are wider than long with a width to length ratio (LP/WP) of 0.50–0.89 (n = 21) (Fig. 35, Table 4). In contrast, the vaginal palpi are broadly rounded with length to width ratio of 0.94–1.36 in the sympatric D. carinulata (Fig. 37) and D. meridionalis (Fig. 38; Table 4). The vaginal palpi are also broadly rounded in D. elongata (Fig. 34). In addition, the width of the widest lobe of the stalk (WLS) of internal sternite VIII (IS VIII) is usually larger in D. carinata (range 0.11–0.17 mm; Fig. 35) compared to D. elongata (range 0.06–0.11 mm; Fig. 34). Some female D. carinata can be distinguished from D. sublineata by having the tips of both lobes (TL) of the stalk of IS VIII strongly curved inward with the tips either pointed or rounded (Fig. 40—Baghdad, Ashgabat, and Pul-e Charki), a combinations of characters never found in D. sublineata (Figs. 36, 41). Some female D. sublineata can be distinguished from D. carinata by having the tips of the lobes of the stalk of IS VIII either not curved or curved outward toward the apical lobe with the tips either rounded or quadrate (Fig. 41—Ndiol, Perpignan, Kom Ombo). Female D. carinata and D. sublineata with at least one lobe of the stalk of IS VIII curved only slightly inward (Figs. 35–36, 40—Lagodekhi and Ardanuc, 41—Biskra and Tamri) are indeterminable to species, except on the basis of geographic distribution outside of the area of known sympatry in Iraq. Measurements. See Tables 2 and 4.

FIGURES 29–33. Palmate (dorsal) endophallic sclerite (dorsal view). 29—Diorhabda elongata, 30—D. carinata, 31—D. sublineata, 32—D. carinulata, 33—D. meridionalis. CES—connecting endophallic sclerite, DS—distal spines, EES—elongate endophallic sclerite, LA—lateral appendage, LN—lateral notch, PES—palmate endophallic sclerite, SDS—subdistal spines. Scale bar 1.0 mm.

Coloration. Subsutural and submarginal elytral vittae are often present in D. carinata (Fig. 9), and the vittae may extend well into the basal half of the elytra, such as in D. sublineata (Fig. 5). In contrast, the elytral vittae, if present, are confined to the apical half of the elytra in D. elongata (Fig. 1). The elytral vittae are less often present in D. carinata (Fig. 3) compared to D. sublineata. Live specimens of D. carinata tend to have

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 59 less greenish-yellow tinting in veins of the elytra (Fig. 4) than do D. elongata (Fig. 2). In contrast, D. sublineata (Fig. 6) and D. carinulata (Fig. 8) both lack greenish-yellow tinting. Type material. According to Berti and Rapilly (1973), the Faldermann type material of D. carinata, consisting of a male holotype, is deposited in the Mniszech Collection at MNHN. We studied the original description by Faldermann (1837), based on an unspecified number of specimens, the illustration of the endophallus of the male holotype by Berti and Rapilly (1973), and topotypes from the Transcaucasus.

FIGURES 34–38. Female genitalia: internal sternite VIII (IS VIII), vaginal palpi (VP), and spermatheca (SP). 34—Diorhabda elongata, 35—D. carinata, 36—D. sublineata, 37—D. carinulata, 38—D. meridionalis. AL—apical lobe of IS VIII, LP—length vaginal palpus, PA—pointed appendage of SP, ST—stalk of IS VIII, TL—tips of lobes of IS VIII, WAL—width apical lobe of IS VIII, WLS—width lobe of stalk of IS VIII, WP—width vaginal palpus, WST—width stalk of IS VIII. Scale bar 1.0 mm.

60 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURES 39–43. Female internal sternite VIII (IS VIII). 39—Diorhabda elongata, 40—D. carinata, 41—D. sublineata, 42—D. carinulata, 43—D. meridionalis. AL—apical lobe, ST—stalk, TL—tips of lobes of stalk, *—IS VIII distinctly identifiable as D. carinata, **—IS VIII distinctly identifiable as D. sublineata. Scale bar 1.0 mm.

Material examined. 122♂♂ dissected (diss.), 67♀♀ diss., 87♂♂, 125♀♀. AFGHANISTAN: 1♂ diss., 3♂♂, Gerab [Sare Gearbid Mt.; 33.89667°N, 66.64583°E], Orurgan [Velayat–e Oruzgan], 1,300 m, 12-VI- 1970, O. Kabakov, ZIN [2004-02] [from series listed as Diorhabda elongata carinata from Uruzgan, Herab by Medvedev (1985)]; 1♂ diss., Polichromi [Pol-e khromi; 35.948889°N, 68.714167°E], 28-V-1956, H.G. Amsel, [from series listed as D. e. carinata by Bechyné (1961)], NHMB [2003-23]; 1♀ diss., Polichromi [Pol–e khromi], 700 m, 5-VI-1956, H.G. Amsel, NHMB [2005-05]; 2♂♂ diss., 1♀ diss., 4♂♂, 6♀♀, Pul–e–Charkhi [Pul–e Charki; 34.541944°N, 69.348889°E], 22–24 km east northeast Kabul, 1,780 m [elev.],

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 61 19-VI-1974, L. Papp, No. 163, D. elongata det. V. Tomov, HNHM [2004-09, 2005-03, 04]; 1♂ diss., Tangi–Gharuh a. Kabul–Fluss [Tangi Gharu Pass, Kabul River; 34.62°N, 69.62°E], 1,600 m [elev.], 8-VII- 1952, J. Klapperich, D. elongata det. K. Lopatin, HNHM [2004-05] [from series listed as D. e. carinata by Lopatin (1963)]; ARMENIA: 1♂ diss., 1♀, Eriwan [Yerevan; 40.181111°N, 44.513611°E], Caucasus, NHMB [2003-39]; 1♂, Eriwan, Caucasus, 1893, Horváth, D. elongata det. K. Lopatin [HNHM]; AZERBAIJAN: 1♂ diss., 2♂♂, 16♀♀, Aresch [a district of the 19th century Russian Caucasus located in present-day northern Azerbaijan according to Frisch (2007)], Caucasus, Schekownikow, D. e. ab. carinata det. Le Moult, IRSNB [2006-08]; 1♀ diss., Dzhafarkhan [39.9431°N, 48.4917°E], 23-V-1939, AMA [2004- 01]; 2♂♂ diss., 2♂♂, 2♀♀, Elisabetpol [Ganca; 40.682778°N, 46.360556E], Caucase, Babadjanides, D. elongata [1♂]; O. Leonhard [collection; 1♂], DEI [2003-08, 2005-09]; 2♂♂ diss., 2♂♂, 2♀♀, Elisabetpol, Caucasus, D. elongata det. K. Lopatin [HNHM], NHMB [2003-53], ZMAN [2008-20], DEI [1♂], HNHM [1♂, 1♀]; 1♂ diss., Geoktapa [Geok–Tapa stream; 39.188056°N, 48.679444°E], 18-VII-1901, P. Schmidt, rice fields, ZIN [2004-22]; 1♀ diss., Geok–Tapa [stream], Transcaucasus, G. C. Champion, BMNH [2003-10]; 1♂ diss., Karandonly [Qaradonlu; 39.7914°N, 48.0428°E], on banks of Araks [Aras] River, 29-V-1911, P. Schmidt, ZIN [2004-04]; 1♂ diss., Khankendy [Xankandi; 39.8153°N, 46.7519°E], Erivansк.g [Erivanskaya gubernia; Xankandi was formerly of Armenia, and now it is in the contested Nagorno-Karabakh zone], Maljushenco, NHMB [2003-42]; 1♂ diss., Mingechaur [Mingacevir; 40.77°N, 47.0489°E], 19-IV-1959, AMA [2004-03]; 1♂ diss., Ordubad [38.9081°N, 46.0278°E], Araxesthal [Aras River], Caucasus, Col[lection] Matcha, NMPC [2004-53]; 1♀, Ordubad, Caucasus, Col[lection] Matcha, NMPC; 4♂♂ diss., 2♂♂, 2♀♀, Ordubat [Ordubad], [19]14, Dr. Veselý, NMPC [2004-10, 55, 2005-38, 39]; 1♂ diss., Ordubat [Ordubad], VI-1910, Javůrek, Coll[ection] Dr. J. Veselý, D. elongata, NMPC [2004-07]; 2♂♂ diss., Qobustan [40.08416°N, 49.41583°E], 18-IV-1984, N. Mirzoeva, AMA [2004-04, 05]; 1♂ diss., Sabirabad [40.0128°N, 48.4789°E], 6-VI-1929, AMA [2004-02]; CHINA: Xinjiang Uygur Zizhiqu:1♂ diss., 1♂, 1♀, Kuldja [Yining; 43.9000°N, 81.35000°E], Tien-Shan, Sven Hedins Exp. Ctr. Asien, D. elongata, NHRS [2008-01]; 1♀ diss., 1♀, Ili [Ili Kazakh Autonomous Prefecture, includes Yining], V-[19]06, Collectie C. & O. Vogt Acq. 1960, ZMAN [2008-18]; GEORGIA: 1♂ diss., 1♀ diss., 1♂, Eldari [41.28833°N, 46.46305°E], 18-IV-1910, G. Vinogradov–Nikítin, ZIN [2004-06, 23]; 1♀ diss., Eldari, 3-VIII-1984, G. Jacobsen, ZIN [2004-03]; 1♂ diss., 1♀ diss., Lagodekhi [41.8228°N, 46.2756°E], 15-V-1910, L. Miokosiewicz, ZIN [2004-09, 24]; 2♂♂ diss., 2♀♀ diss., 2♂♂, Owtshaly [Avchala; 41.79028°N, 44.8261°E], 24-VI-1879, G. Sievers, ZIN [2004-08, 11, 25, 26]; 1♂ diss., 1♂, 1♀, Mtzchet [Mts'khet'a; 41.8439°N, 44.7164°E], Caucasus, [18]79, Leder (Reitter), ZIN [1♂ diss., 2004-19], NMPC; 1♂ diss., 1♀, Tflis [T’blisi; 41.725°N, 44.79083°E], Caucasus, Leder (Reitter), D. elongata, Coll. Reitter, HNHM [2004-01], NMPC; IRAN: 2♂♂ diss., 2♀♀ diss., Borazjan, 30 km north northeast [29.53333°N, 51.40000°E; 10 km N Dalaki, Hableh Rud River; Tamarix sp. present (Hoberlandt 1983)], 18-19-IV-1977, Exped. Nat. Mus. Praha Loc. No. 298, NMPC [2004-41, 48, 2005-36], MPBF [2003-04]; 2♂♂ diss., Jeiugir (33° 27' N, 49° 01' E) [sic; Jolow Gir; 32.96667°N, 47.81611°E; coordinates corrected from relation to Sarab–e Jahangir], 20 km southeast Sarab–e Jahángir [32.326994°N, 48.51111°E], Lorestán, 8-10-X-1998, Chvojka, D. elongata det. A. Warchalowski, NMPC [2004-11, 2006-02]; 2♂♂ diss., 4♀♀, Omidiyeh, 34 km south southeast [30.55°N, 49.91667°E; Rud-e Zohreh river, Tamarix sp. present (Hoberlandt 1983)], 16-17-IV-1977, Exped. Nat. Mus. Praha Loc. No. 292, NMPC [2004-33, 2005-34]; 1♂ diss., 1♂, Golhak [Qolhak; 35.7803°N, 51.4919°E], 1400m [elev.], near Teheran [Tehran], VI-VIII, 1961, J. Klapperich, D. elongata det. I.K. Lopatin, HNHM [2003-02]; 1♀ diss., Robat–e Qozlog [Robate–Ghozlog; 36.7°N, 54.61667°E; valley of small river; Tamarix not recorded (Hoberlandt 1974)], [10 km] south of Gorgan, 500 m [elev.], 26-VII-1970, Exped. Nat. Mus. Praha Loc. No. 74, D. elongata det. I.K. Lopatin 1964, NMPC [2004-04] [from series listed as D. elongata by Lopatin (1981)]; 1♂ diss., 1♀, Semnan to Dameqan [Damghan] road, 35° 46' 13" N, 53°, 44' 47" E [35.77028°N, 53.74639°E], 1,975 m elev., 13-V-2000, J. Gaskin, on Tamarix, No. 870, GSWRL [2003-38]; 1♂ diss., 1♂, 3♀♀, Shushtar [32.05°N, 48.85°E; river Karun; Tamarix sp. present (Hoberlandt 1983)], 13-IV-1977, Exped. Nat. Mus. Praha Loc. No. 287, NMPC [2004-35a]; 2♀♀ diss., Wildlife Park, vicinity of Dasht, 650 m [elev.] [37.38°N, 55.89°E; Mohammad Reza Shah Wildlife Park (= Golestan Biosphere Reserve), east tributary of the river

62 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Rud-i Gorgan; Tamarix not recorded; coordinates adjusted based on elevation and forest description (Hoberlandt 1974)], 27-30-VII-1970, Exped. Nat. Mus. Praha Loc. No. 77, NMPC [2004-17, 20]; IRAQ: 4♂♂ diss., 5♀♀ diss., 4♀♀, Assur [Ash Sharqat; 35.45861°N, 43.25722°E], , 1910, Pietschmann, D. elongata [1♀ diss.], Mesopot. Exp. Nat. O.V. 1910, NHMB [2003-06, 21, 33, 34, 35, 36, 47, 51, 2004-04] [from series listed as D. elongata by Holdhaus (1920)]; 1♂ diss., Assur [Ash Sharqat] Mesopotamia, Coll. F. Hauser, NHMB [2006-01]; 1♂ diss., Baghdad [33.33861°N, 44.37722°E], summer 1923, B.W. G. Hingston, B.M. 1923-486, BMNH [2005-03]; 2♂♂ diss., 2♀♀ diss., 3♀♀, Baghdad, IV-1936, Frey, NHMB [2003-07, 22, 32, 50]; 4♂♂ diss., 1♀ diss., 5♀♀, 4♂♂, Baghdad, Kálalová, NMPC [2004-06, 09, 13, 54, 56]; 1♀ diss., Khanikin [Khaniqin; 34.34694°N, 45.40056°E], 9-IV-1936, Frey, D. persica Faldermann det. H. Bollow 1938, NHMB [2003-37]; KAZAKHSTAN: 1♀ diss., Aulie–Ata [Taraz; 42.9°N, 71.3667°E], Turkestan, K. Arris, D. sulphureus Reitter det. H. Bollow 1939, D. e. subsp. carinata Fald. det. J. Bechyné 1955, NHMB [2003-30]; 2♀♀, Aulie Ata, Syr Darja, NMPC; 1♂ diss., Chilik [Shelek] near, Chilik River Valley [43.6003°N, 78.2963°E], 6-VI-1999, I.D. Mityaev and R. Jashenko, on Tamarix ramosissima, voucher shipment GSWRL-1999-8, GSWRL [2004-01] [28 specimens from Lot GSWRL-1999-1 identified as D. elongata by A. Konstantinov on 20-VII-1999 at USDA Systematic Entomology Laboratory (SEL), SEL Lot # 9904498]; 2♂♂ diss., Chundzha Village, Charin [Charyn] River [43.5161°N, 79.2437°E], Gorodinski, V-1994, D. elongata det. D. Sassi 2000, GSWRL [2003-58, 59]; 1♂ diss., Sargotay [Sarytogay] Locality [43.43240°N, 79.15040°E], Charyn River, 700 m [elev.], 2-VII-1907, A.G. Jacobson, ZIN [2005-01]; 1♀ diss., Shelek, near, 1-VIII-1999, I.D. Mityaev and R. Jashenko, on Tamarix ramosissima, voucher shipment GSWRL-1999-15, GSWRL [2005-94]; KYRGYZSTAN: 1♂ diss., 2♂♂, Kugart–su River [Kugart stream], Ferghana Valley [40.8667°N, 72.8833°E], 8-V-1925, F. Dobrzhanskii, ZIN [2004-10]; PAKISTAN: 1♀ diss., Kund [33.9306°N, 72.2289°E], 4-VI-1976, CIBC [Commonwealth Institute of Biological Control], larvae feeding on T. aphylla, D. elongata, Tam-6/76-147-II, 2364, C.I.E. coll. A 9076, CABP [2003-01] [from series listed as D. elongata by Habib and Hasan (1982)]; 1♀ diss., Nomal [36.07305°N, 74.25444°E], 25-VI-1962, CIBC, adult feeding on T. cf. indica [as T. cf. troupii], D. elongata, CIBC Wol-6/62-465-III, 947, CIE. Coll. No. 18503, CABP [2003-02]; 1♂ diss., 1♀ diss., Shorkot [31.90972°N, 70.87722°E], west Pakistan, 17-IV- 1967, M. Nararullah, on Tamarix sp., D. elongata N.A. Aslam det. 1970, D. elongata det. M.L. Cox 1982, CIE Coll. No. 3509, Comm. Inst. Ent. B.M. 1981-315, BMNH [2003-06, 07]; SYRIA: 2♀♀ diss., 1♀, Alep[po] [Halab; 36.20278°N, 37.15861°E], Syrie, NHMB [2003-14, 44]; TAJIKISTAN: 1♀ diss., 1♀, Dushanbe [38.5600°N, 68.7739°E], May–June 1966, Král, NMPC [2005-32]; 1♂ diss., 2♂♂, 1♀, Khodzhent [Qayraqqum; 40.2647°N, 69.7894°E], Ob[last], 23-IV-1903, Val'nev, ZIN [2004-13]; 1♂ diss., 3♂♂, 1♀, Ghissar and Karateghin Mt. Ranges [approximate location; 39.06°N, 69.82°E], ZIN [2004-27]; 1♂ diss., Hissar [Hisor; 38.48194°N, 68.59694°E], Buchara, [D.] sulphurea, MSNM [2003-16]; 1♂ diss., 1♂, Karatack [Karatag; 38.617°N, 68.333°E], Buchara, Stauding, [D.] sulphurea m.n.sp., D. sulphurea Reitt., DEI [2003-11]; 2♂♂ diss., 4♂♂, 4♀♀, Karatag, west Buchara, 916 m [elev.], 1898, F. Hauser, D. [e.] a. carinata [2♂♂ diss., 1♂, 1♀ of DEI], DEI [2005-12], NHMB [2005-07; 3♂♂, 2♀♀ not diss.], BMNH [1♀]; 1♀ diss., Karatag, west Buchara, D. [e.] v. carinata, NMPC [2005-33]; 1♀ diss., Kurgan–Tsu BG [Qurghonteppa or Kurgan–Tyube; 37.83639°N, 68.78028°E], Ruβland, 28-IV-1989, D. elongata R. Beenen det. 2002, RBCN [2003-09]; 2♂♂ diss., 1♀, Nurek [Norak; 38.38833°N, 69.325°E], Dušanbe [Dushanbe] environs, 25-VI- 1976, Josef Král, NMPC [2004-38, 39]; 2♂♂, 2♀♀, diss., Sary-pul [38.4167°N, 70.1333°E], Mts. Karategin, 1,482 m [elev.], 1898, F. Hauser, D. e. v. carinata R. det [1♂ of BMNH], NHMB [2003-29], BMNH [2005-04 and 1♀]; 1♂ diss., Vachsch [Vakhsh] River [approximate location: 37.5629°N, 68.5275°E], 23-IV-1992, RBCN [2003-15]; TURKEY: 1♀ diss., 3♂♂, 7♀♀, Ağri [Agri], 9 km west on road to Horasan [39.7393°N, 42.9235°E], 1,600 m [elev.], Ağri Prov., 1-VIII-2000, C. Morkel, on Asteraceae, D. elongata det. D. Erber 2001, ZMHB [2006-01]; 1♂ diss., Aralich [Aralik; 39.872778°N, 44.519167°E], Caucasus, 1893, Horváth, [D.] e. var. sublineata det. K. Lopatin 1958, [HNHM 2003-09]; 1♂ diss., 1♀ diss., Ardanuç [Ardanuc; 41.12861°N, 42.05916°E], 600m, Artvin [Prov.], 9-VII-1998, I. Aslan, D. elongata det. I. Aslan, IAET [2003- 01, 02] [from series listed as D. elongata by Aslan et al. 2000]; 1♂ diss., 2♀♀, Artvin [41.18222°N, 41.81944°E], 2-VII-1975, 700 m [elev.], Osella, MRSN [2003-06]; 1♂ diss., 1♂, 1♀, Artvin, 13–15-VIII-

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 63 1976, Bohao, ex coll. Josef Kral, Prag, 1987–1989, coll. U. Arnold, Berlin, D. elongata carinata R. Beenen det 1993, RBCN [2008-01]; 1♀ diss., Artvin, 12 km east [41.1329°N, 41.8978°E], 26-V-1990, P. Kanaar, D. elongata ssp. carinata R. Beenen det. 1992, RBCN [2003-06]; 1♀ diss., 6♀♀, Siirt, southwest, on riverbank [37.8192°N, 41.8679°E], ca. 550 m [elev.], southeast Anatolia, 28-VII-1985, Heinz, D. e. carinata det. Beenen 1996 [2♀♀], D. elongata det. Beenen 1998 [1♀ diss., 4♀♀], ZMHB [2006-04]; TURKMENISTAN: 2♂♂ diss., 1♀ diss., Ashgabat, 2-IX-2002, S.N. Myartseva, on T. ramosissima, GSWRL [2003-02, 04, 07]; 2♂♂ diss., Ashgabat [37.95°N, 58.36667°E], Dry Sport Lake, 26-IX-1996, S. N. Myartseva, on Tamarix sp. [prob. T. aralensis J. Gaskin], [parasitoid] wasp found in abdomen [1♂], GSWRL [2005-36, 102] [7 specimens from Lot GSWRL(CJD)-1996-4 identified as D. elongata by S.M. Clark on 28-X-1996 at West Virginia Dept of Agric.] [8 specimens (no. 9204) from Lot GSWRL(CJD)-1997-1 identified as Diorhabda n. sp. by A. Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023] [parasite Baryscapus diorhabdivorus Gates & Myartseva (Hymenoptera: Eulophidae) found in 1 adult in lot GSWRL-2005-01 identified by M. Gates on 28-VII-2005 at USDA SEL]; 1♀ diss., 4♂♂, 7♀♀, Ashgabat, Dry Sport Lake, 11- V-1997, S. N. Myartseva, on T. ramosissima [prob. T. aralensis J. Gaskin], voucher shipment GSWRL(CJD)- 1997-11, 9005–9012, 9020–9022, D. elongata det. I. Lopatin 1999 [4♂♂, 7♀♀], GSWRL [2005-66] [8 specimens (nos. 9200, 9201) from Lot GSWRL(CJD)-1997-1 identified as Diorhabda n. sp. by A, Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023]; 4♂♂ diss., 2♀♀ diss., Ashgabat, Dry Sport Lake, 11-VI-1997, S. N. Myartseva and P. Boldt, on T. ramosissima [prob. T. aralensis J. Gaskin], USDA lab colony at Temple, Texas, voucher (1997, J.L. Tracy), voucher shipment GSWRL(CJD)-1997-12, D. elongata det. I. Lopatin 1999 [1♂ diss.], 9004, GSWRL [2003-11, 2005-09, 59, 60, 61, 62]; 7♂♂ diss., 7♂♂, 5♀♀, Ashgabat, Dry Sport Lake, 17-18-VIII-1998, A Knutson, Andrey Averin and Allen Knutson [7♂♂], on T. ramosissima [prob. T. aralensis J. Gaskin], defoliating shrubs, voucher shipment GSWRL-1998-15, D. elongata det. I. Lopatin 1999 [1♂ diss., 5♂♂, 3♀♀], 8993–8996, 8998–9003, GSWRL [2005-25; 2008-12, 13, 14, 15, 16, 17] [photo of damage to tamarisk in DeLoach et al. (2003b), Fig. 3]; 1♀ diss., 6♂♂, 2♀♀, Ashgabat, Dry Sport Lake, 26-IX-2002, S.N. Myartseva, on Tamarix spp., GSWRL [2006-10]; 1♂ diss., 3♀♀ diss., Ashgabat, Karakum [Quaragum] Canal, 12-IX-2002, S.N. Myartseva, on T. ramosissima, GSWRL [2003-A1, A2, A3, A4]; 2♀♀ diss., Cardruj [Turkmenabat; 39.10139°N, 63.575°E], Buchara, [D.] e. var. sublineata det. K. Lopatin 1958, HNHM [2003-13, 2004-02]; 1♂ diss., 2♂♂, Chardzhuy [Turkmenabat], 21- VI-[19]04, Z. Fisher, ZIN [2005-03]; 2♂♂ diss., Cemenibit [35.4489°N, 62.3958°E], 22 km north Kujka, 1- IV-1992, Snilek, EGRC [2005-02, 07]; 1♀ diss., Chule [Chuli] Canyon, 53 km west Ashgabat (10, 852 km odom.) [38.08°N, 57.77°E], 25-V-1995, on Tamarix ramosissima, C.J. DeLoach, GSWRL [2004-13]; 1♂ diss., Danata Village [39.08389°N, 55.1617°E], 157 m [elev.], 31-V-2000, J. Gaskin, on T. aucheriana [det. J. Gaskin], Coll. No. 1044, GSWRL [2003-55]; 3♂♂ diss., 3 ♀♀ diss., Svintsovyi Rudnik Village [Svintsovyy Rudnik; 37.88472°N, 66.45138°E], Kughitangolarja [Kugitang] River, Kughitang Mts., 3-V-1989, Osipov Coll., GSWRL [2003-66, 67, 68, 69, 2004-06, 07]; 1♂ diss., 1♀ diss., 1♂, 1♀, Tedzhen [Tejen; 37.37861°N, 60.49611°E], D. elongata, Coll. Reitter, HNHM [2004-03, 04]; 1♀ diss., 1♀, Tschardsui [Turkmenabat; see Cardruj], Turkestan, D. sulfurea Rt. det. Dr. Fleischer, NMPC [2005-03], 1♀, Tschardsui [Turkmenabat], Trans-Caspia, 4358b, coll[ection] Kǒuřil P5/46/62, NMPC; UKRAINE: 1♂ diss., 1♀ diss., Izium v[illage] [geocoordinates not locatable], Odessa Reg. [mapped approximate location in Odessa region as Izmayil: 45.3500°N, 28.8330°E], southwest Ukraine, 15-VII-1974, Osipov Coll., GSWRL [2003-71, 2004-09]; UNITED STATES OF AMERICA (introduced): Texas: Hutchinson Co.: 4♂♂ diss., 1♀, Johnson Ranch (cage 7) on Canadian River near Borger [approx., 35.7294°N, -101.4126°W], 26-VI-2007, E. Jones, on T. ramosissima/T. chinensis, voucher [source: 7 km W of Qarshi, Uzbekistan], GSWRL [2007-54, 55, 56, 57]; Baylor Co.: 1♂ diss., 2♂♂, 1♀, Lake Kemp, west side, Waggoner Ranch site B, 33.709267°N, - 99.276395°E, 7–9-X-2008, Charles Randal, on T. ramosissima/T. chinensis, voucher [source: 7 km W of Qarshi, Uzbekistan], GSWRL [2008-25]; UNKNOWN COUNTRY: 1♂ diss., Attica [location of Attiki, Greece; considered as mislabeled], Reitter, ZMAN [2008-05]; 1♂ diss., 1♀, Araxes [Aras River of Armenia, Azerbaijan and Iran], VI-[19]10, J. Vesely, Coll[ection] Dr. J. Veselý, NMPC [2004-23]; 2♂♂, 2♀♀, Araxes, coll. Purkynĕ, ZMAN; 1♀ diss., Araxesthal [Aras River], Reitter, HNHM [2003-12]; 1♂ diss., Dulicata

64 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS [geocoordinates not locatable, possibly Dul'ta, Uzbekistan], Turkestan, NHMB [2003-03]; 1♂ diss., Ferolash [geocoordinates not locatable], Caucasus, D. elongata, DEI [2005-10]; UZBEKISTAN: 2♂♂ diss., 3♀♀ diss., Buxoro, 5 km northwest [39.8228°N, 64.39134°E], along the road to Gazli, 27-IX-2002, R. Sobhian, on Tamarix, USDA [Buchara] lab colony at Albany, California, voucher (2002-2003, D. Bean), shipment EIWRU-2002-1010, GSWRL [2003-39, 2005-52, 55, 57, 58]; 1♂ diss., 1♀ Ferghana [Farg`ona; 40.3933°N, 71.7794°E], V-1961, L. Medvedev, D. elongata det. L. Medvedev, GSWRL [2004-12]; 3♂♂ diss., 3♀♀ diss., 6♂♂, 4♀♀, Karshi [Qarshi], 10 km west [38.92278°N, 65.73307°E], 7-IV-1999, A. Kirk & R. Sobhian, on Tamarix, D. elongata det. I.K. Lopatin 1999 [3♂♂ diss., 3♀♀ diss., 4♂♂, 3♀♀], 9052–9064, GSWRL [2003- 44, 2005-71, 72, 2007-02, 2008-18, 2008-19]; 4♂♂ diss., 5♀♀ diss., Qarshi, 7 km west [38.90721°N, 65.76314°E], 26-IX-2002, R. Sobhian, on Tamarix, USDA lab colony at Temple, Texas, voucher (2003, L. Milbrath [nos. 1298-1300]), USDA lab colony at Albany, California, voucher (1♂, 3♀♀, 26-VI-2002 and 20- I-2003, D. Bean), shipment EIWRU-2002-1010, GSWRL [2003-16, 18, 72, 73, 2005-69, 70, 85, 88, 89] [released in north Texas in 2006] [used in biological studies of Milbrath and DeLoach (2006a, 2006b), Milbrath et al. (2007), Herr et al. (in prep.), Bean and Keller (in prep.), and Thompson et al. (in prep.)]; 1♂ diss., 2♂♂, Namangan City [40.99527°N, 71.6725°E], Ferghana Valley, 30-III to 15-IV-1903, Yankovskii, ZIN [2004-07]; 1♀ diss., 4♀♀, Samarkand [Samarqand; 39.6542°N, 66.9597°E], Turkestan or[iental], daroval Král, NMPC [2006-03]; 1♂ diss., Sansar [Kirk; 39.7167°N, 68.0167°E], Turkestan, 1892, Glasnov, NHMB [2004-02]; 2♂♂ diss., 4♀♀, 2♂♂, Saratoff. [Saratovskiy; 40.7667°N, 68.6667°E], Turkestan, NHMB [2003-24, 2005-06]; 2♂♂ diss., 1 ♀ diss., 6♂♂, 5♀♀, Tashkent [Toshkent], 140 km south [approximate location: 40.29057°N, 68.91332°E], 10-IV-1999, Kirk & Sobhian, on Tamarix spp., D. elongata det. I. Lopatin 1999 [1♂ diss., 1♀ diss., 6♂♂, 3♀♀], 9067–9077, GSWRL [2003-51, 52, 2004-16]; 1♂ diss., 7♂♂, 9♀♀, Tashkent [Toshkent], 50 km east [41.3206°N, 69.8273°E], 11-IV-1999, Kirk & Sobhian, on Tamarix spp., D. elongata det I. Lopatin 1999 [6♂♂, 6♀♀], 9078, 9080–9090, GSWRL [2003-81]. Distribution. General. The native distribution of D. carinata ranges from Ukraine, eastern Turkey and Syria east to northwest China, Kyrgyzstan and Pakistan, extending as far south as southern Iran (Maps 3, 6). Further collections could give an exact location for D. carinata within the Odes'ka Oblast' of Ukraine and possibly expand its range to include the coast of the Black Sea in eastern Romania and southern Russia, and the semi-arid regions of northwestern India. Confirmed Records. We have dissected specimens listed above (see Materials Examined) and confirmed the presence of D. carinata in the following countries with previous literature records (Map 3): Armenia (as Transcaucasus, Faldermann 1837), Azerbaijan (as Transcaucasus, Faldermann 1837), Georgia (as Transcaucasus, Faldermann 1937), Turkmenistan (as D. e. var. carinata) (Jacobson 1901), Afghanistan (as D. e. carinata) (Bechyné 1961; Lopatin 1963; Wilcox 1971; Medvedev 1983, 1985), Kazakhstan, and China (as D. e. carinata) (Lopatin et al. 2004). Specimens were dissected from three of four regions of Afghanistan listed with D. e. carinata by Medvedev (1983) (central, Badakshan, and Kabul-Jalalabad) and close to the fourth region (western) at Cemenibit, Turkmenistan. Diorhabda carinata were dissected from three series identified in publications as D. e. carinata in Afghanistan: Polichromi [Pol–e khromi] (Bechyné 1961), Tangi–Gharuh [Tangi Gharu Pass] at Kabul River (Lopatin 1963), and Gerab Orugan [Sare Gearbid Mt, Velayat–e Oruzgan] (Medvedev 1985). New Records. We have dissected D. carinata from the following countries for which we find no previous specific reports of D. carinata in the literature: Turkey, Iran, Iraq, Syria, Uzbekistan, Kyrgyzstan, Tajikistan, Pakistan, Ukraine, and the United States of America (Texas; introduced; Map 7, see Potential in Tamarisk Biological Control below for additional details). With the exception of Ukraine and Kyrgyzstan, past reports of D. elongata occur from all these countries that should refer, at least in part, to D. carinata (see synonymy above). Yakhontov and Davletshina (1955) site a potential report of D. elongata in Ukraine by Degtarev (1928), but we were unable to obtain this reference for verification. We dissected D. carinata from four series with published identifications of D. elongata: Assur [Ash Sharqat], Iraq (Holdhaus 1920); Robat–e Qozlog [Robate–Ghozlog], Iran (Lopatin 1981); Kund, Pakistan (Habib and Hasan 1982); and Artvin, Turkey (Aslan et al. 2000).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 65 Unconfirmed Records. We cannot confirm a listing of D. carinata (as D. e. var. carinata) from Russia (Heyden 1891), but suspect it may be present along the Caspian Sea in the Dagestan region of southern Russia (Map 8b). We consider records of D. e. var. carinata from France (Laboissière 1934) and D. e. ab. carinata from Italy (Porta 1934) as erroneous (see lists of material examined with these identifications by Laboissière for specimens from France and Italy under D. sublineata and D. elongata, respectively). We consider as mislabeled a single male specimen bearing the label of “Attica, Reitter” (Greece) from ZMAN (2008-05). We dissected only D. carinata from all seven available locations in Afghanistan (four) and Pakistan (three) (Map 3). Therefore we consider as D. carinata five unconfirmed records of D. elongata from northern Pakistan (Habib and Hasan 1982; three records) and D. e. carinata from Afghanistan (Medvedev 1985; two records), but D. carinulata may also be present at some locations. Specimens of D. carinata were dissected from all nine available localities in Azerbaijan and D. carinulata was also dissected from two of these same localities. Consequently, we accept all nine unconfirmed collection records of D. elongata in Azerbaijan (Samedov and Mirzoeva 1985) as D. carinata, but D. carinulata may also be present at some of these localities. Mirzoeva (2001) reported that D. carinata (as D. elongata) was found in all four major regions of Azerbaijan and was most common in the Great Caucasus and Small Caucasus. Specimens dissected from all five available locations in eastern Georgia were D. carinata, and we consider Lozovoi’s (1961) report of D. elongata from eastern Georgia to be D. carinata. We dissected D. carinata from all five available locations in Turkey east of 41°E, including three locations in Arvin Ili. The closest occurrence of D. elongata to D. carinata in Turkey is in Malatya, which is 316 km north northwest of D. carinata in Siirt. Therefore, we regard two unconfirmed records of D. elongata in Artvin Ili and neighboring Ezurum Ili to also be D. carinata. We dissected D. carinata from nine and D. carinulata from one of the ten available locations in Tajikistan. Therefore we consider nine unconfirmed records in Tajikistan to be D. carinata, but D. carinulata may also be present at some sites. We regard a report of D. elongata at Mikhaylovka [Sarykemer], Kazakhstan (Kulenova 1968) to be D. carinata because of its proximity to a collection of D. carinata from Auliye–Ata [Taraz], Kazakhstan. Below are 28 unconfirmed locality records that we consider as D. carinata, but are listed as D. e. carinata in Afghanistan (2) and D. elongata in Azerbaijan (9), Kazakhstan (1), Pakistan (3), Tajikistan (9), and western Turkey (4) (Map 3): AFGHANISTAN: 2 spms., Banu [Banow], southwest [35.63420°N, 69.25260°N], Baglan [Velyat–e Baghlan], 2,000 m [elev.], 14-VIII-1972; 1 spm., Samti [west; 37.5854°N, 69.9592°N], Pan’ [Panj] River, Badakhshan [Velyat–e Badakhshan], 1,100 m [elev.], IV-1971 (Medvedev 1985); AZERBAIJAN: Akstafa [Agstafa; 41.1189°N, 45.4539°E], 25-V-1958; Belokany [Balakan; 41.7258°N, 46.4083°E], 3-V-1972; Dzhul'fa [Culfa; 38.9500°N, 45.6319°E], 13-V-1933; Matsekh [Matsex; 41.6553°N, 46.5811°E], 3-V-1972; Mingachaur [Mingacevir; 40.7700°N, 47.0489°E], 23-V-1946 [D. carinata dissected from same location]; Neftechala [Neftcala; 39.3586°N, 49.2469°E], 28-V-1980; Saatly [two possible geocoordinates], 26-VI-1969; Sumgait [Sumqayit; 40.5897°N, 49.6689°E], 12-VII-1935; Yevlakh [Yevlax; 40.6172°N, 47.1500°E], 12-VI- 1903 (Samedov and Mirzoeva 1985); KAZAKHSTAN: Mikhaylovka [Sarykemer; 43.0000°N, 71.5000°E] (Kulenova 1968); PAKISTAN: Chitral [35.8419°N, 71.7819°E], Tamarix spp.; Gilgit [35.9167°N, 74.3000°E], Tamarix spp.; Islamabad [33.7000°N, 73.1667°E], Tamarix spp. (Habib and Hasan 1982); TAJIKISTAN: Baljuan [Baljuvon; 38.3108°N, 69.6669°E], 11-V-1951, E.P. Luppova; Dushanbe [38.5600°N, 68.7739°E] [D. carinata dissected from same location], P.N. Kulinich; Kal'aiKhumb, behind and on right side, on Pyanj River [Darya–ye Panj] [38.4489°N, 70.81529°E], 6-VII-1960, L.V. Soboleva; Khoja Obigarm [38.9003°N, 68.8008°E], P.N. Kulinich; Kondara [38.1190°N, 68.8283°E] canyon, VI-VII-1956, P.N. Kulinich (Kulinich 1962); Kurgan–Tyube [Qurghonteppa; 37.8364°N, 68.7803°E] environs [D. carinata dissected from same location] (Lopatin 1959); Pyanj [Panj; 37.2383°N, 69.0969°E] environs, P.N. Kulinich (Kulinich 1962); Shaartuz [Shahrtuz; 37.2594°N, 68.1347°E] (Lopatin 1959); Vaidar village [Vaydara; 38.9667°N, 70.1833°E], 20-V-1961, L.V. Soboleva (Kulinich 1962); TURKEY: Artvin [41.1822°N, 41.8194°E], 5-VII-1994, I. Aslan [D. carinata dissected from same location]; Sarigol [40.9553°N, 41.4989°E],

66 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS 5-VII-1994, I. Aslan; Yusufeli [40.8167°N, 41.5500°E], 5-VII-1994, I. Aslan (Aslan et al. 2000); 2♂♂, 3♀♀, Uzundere [40.5314°N, 41.5531°E], Ezurum Ili, 1,100 m [elev.], 7-VI-1995 (Aslan 1998). Discussion. Taxonomy. Galeruca carinata Faldermann (1837) (as Galleruca carinata) was described from the Transcaucasus and synonymyzed under G. elongata by Reiche and Saulcy (1858). Weise (1893) proposed the variety D. elongata var. carinata under which he incorrectly synonymyzed G. carinulata (Desbrochers 1870) of southern Russia. Weise (1924) later proposed the aberration D. elongata ab. carinata and this has been followed by several taxonomists (Winkler 1924–1932, Laboissière 1934, Warchalowski 2003). Bechyné (1961) proposed the subspecies D. e. carinata with an implied range from the Transcaucasus to central Asia and Afghanistan. Several other taxonomists followed Bechyné in reporting D. e. carinata from Afghanistan (Lopatin 1963, Wilcox 1971, Medvedev 1983), Kazakhstan, and China (Lopatin et al. 2004). Berti and Rapilly (1973) studied the endophalli of type specimens of D. carinata and D. carinulata and restored their species status, removing them from synonymy with one another and, by implication, with D. elongata. However, Berti and Rapilly (1973) omit D. elongata in their discussion and provide no distributional data for D. carinata. We find that the external characters they provide regarding elytral carinae and the shape of the pronotum are variable and insufficient for species diagnosis (further discussed below). In addition, they provide no information on variability of the endophallus. This lack of data has apparently contributed to the lack of recognition of D. carinata as a species in recent taxonomic treatments (Riley et al. 2003, Warchalowski 2003, Bieńkowski 2004, Lopatin et al. 2004). We dissected 25 males from 16 locations of the type locality of D. carinata in the Transcaucasus (Georgia, Armenia and Azerbaijan) with endophalli matching that of the type specimen of D. carinata as illustrated by Berti and Rapilly (1973) (Map 3). One male and two female D. carinulata were dissected from two of the same locations with D. carinata in Azerbaijan. Three key characters of the endophallus of D. carinata that distinguish it from other members of the D. elongata group can be seen in the illustration of the holotype (Fig. 18 of Berti and Rapilly 1973) and our illustration (Fig. 15): (1) the presence of spines irregularly spaced along the distal blade of the elongate endophallic sclerite; (2) a lateral appendage on the palmate endophallic sclerite; and (3) the lack of a connecting endophallic sclerite between the palmate and elongate sclerites. We are certain that specimens we studied with endophalli matching that of D. carinata (Figs. 15, 20, 25, 30) form a single species conspecific with D. carinata. We find additional characters in the endophallic sclerites and female genitalia (vaginal palpi and internal sternite VIII) of D. carinata throughout its range from west to central Asia that distinguish it from D. elongata, D. carinulata and other members of the D. elongata group (Figs. 15, 20, 25, 30, 35, 40; Map 3). The distinctive genitalic characters of D. carinata are maintained in the same areas where D. elongata, D. carinulata, D. meridionalis, and D. sublineata occur, and this is strong evidence for reproductive isolation between these species (see Biogeography below; Map 1, Table 8). Therefore, we firmly support Berti and Rapilly in restoring D. carinata to species status, removing it from synonymy with both D. elongata and D. carinulata. If D. carinata were an interbreeding subspecies of D. elongata, we should see intermediate morphologies of the distinguishing characters found in the two endophallic sclerites and the vaginal palpi. Such intermediate forms should increase along a geographic gradient approaching range contact of these species in Turkey, Georgia and Syria. For example, we should see intermediate forms of the palmate sclerite in a progression from broadly rounded (in D. elongata; Fig. 29) to truncate serrate (in D. carinata; Fig. 30). However, no intermediate forms of palmate sclerites were seen in field collections of 157 male D. elongata and 122 male D. carinata, including 21 male D. elongata in Turkey and Syria and 11 male D. carinata in Turkey and Georgia. Similarly, intermediate forms in vaginal palpi ranging from broadly rounded (as in D. elongata; Fig. 34) to triangulate (as in D. carinata; Fig. 35) were not seen in 85 female D. elongata or 67 female D. carinata, including 14 female D. elongata from Turkey and 9 female D. carinata from Turkey, Georgia, and Syria (Map 1). The lack of intermediate forms is evidence of reproductive isolation between D. elongata and D. carinata. Further evidence for reproductive isolation between D. carinata and several members of the D. elongata group is also found in previously discussed differences in component ratios of putative aggregation pheromones and reduced hybrid egg viability.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 67 Faldermann (1837) gave a length of 7 mm for the male type of D. carinata which is too large to have included the sympatric D. carinulata in which males only reach 6.09 mm (Table 2). Male D. carinata commonly exceed 7 mm in length (range 5.12–7.34 mm). The application of the name “D. elongata” to both D. carinata and D. carinulata where these species are sympatric can be seen in the given range in length of 4.5–8 mm for males and females of “D. elongata” in taxonomic keys of Russia (Ogloblin 1936) and central Asia (Medvedev 1959, Lopatin 1977b, Lopatin and Kulenova 1986). Overall ranges in length for both sexes are 4.63–6.99 mm for D. carinulata and 5.05–8.44 mm for D. carinata (Table 2). We find that external characters used in separating D. carinata from sibling species are inadequate and that genitalic characters must be used. Weise (1893) noted black markings on the pronotum and abdominal sternites as characters distinguishing D. carinata (as D. e. var. carinata) from D. elongata, but this character is highly variable in both taxa. Black spots on the pronotum were also noted as a distinguishing character for D. carinata (as D. e. ab. carinata) by Laboissière (1934) and Porta (1934), who also noted black spots on the femora of the leg. Warchalowski (2003) notes black spots on both the pronotum and head as distinguishing characters for D. carinata (as D. e. ab. carinata). However, we find that black spots on the head, pronotum, and femora variably occur in both D. elongata and D. carinata, making these characters unsuitable for species diagnosis. Bechyné (1961) noted two differences between D. e. carinata and the Mediterranean form, D. e. elongata: (1) D. carinata reaches a greater length, ca. 7 mm and (2) the sublateral carina approaches the lateral margins of the elytra more closely towards the apex than in D. e. elongata. However, both of these external characters are variable and overlap between D. carinata and D. elongata (although it is uncommon for D. elongata to exceed 7 mm in length). We have dissected specimens of D. carinata that were misidentified by taxonomists using external diagnostic characters as D. elongata in eleven countries and D. e. var. sublineata in Turkey and Turkmenistan (see Material examined). Berti and Rapilly (1973) proposed that the following five external characters of D. carinata can be used to distinguish it from D. carinulata: (1) length of ca. 6.7 mm, (2) posterior angles on the pronotum are more pronounced and farther from the base, (3) pronotum is broader, (4) only a lateral carina is clearly present on the elytra, and (5) punctuation on the elytra is simple. They give the following contrasting characters for D. carinulata: (1) length of ca. 4.9 mm, (2) posterior angles on pronotum are less pronounced and closer to the base, (3) pronotum is narrower, (4) in addition to a lateral carina on the elytra, a median carina (along a clear humeral furrow) and a sutural carina are seen, and (5) punctation on the elytra is almost confluent in places. In keys for separating D. carinulata from D. elongata (considered as including D. carinata), Warchalowski (2003) used three of the above characters: body length, the posterior angle of the pronotum, and number of elytral carinae. However, we find all these external characters as too interspecifically variable for species diagnosis, and the only taxonomically reliable characters provided by Berti and Rapilly (1973) are their well illustrated differences in the morphology of the endophalli of D. carinata, D. carinulata, and D. meridionalis. We consider D. e. carinata as regarded by Bechyné to be synonymous with D. carinata. Because of the incorrect synonymization of D. carinulata (as G. carinulata) under D. elongata var. carinata by Weise (1893), Wilcox (1971) included D. carinulata under the name D. e. carinata. But Bechyné (1961) did not mention D. carinulata in his description of D. e. carinata and the larger stated size of ca. 7 mm would exclude almost all D. carinulata (Table 2). We examined a specimen of D. carinata from the series that Bechyné referred to D. e. carinata from Pol–e khomri (as Polichromi), Afghanistan. Also, a specimen of D. carinata was dissected from Taraz (= Auliye–Ata), Kazakhstan with a determination label of D. e. carinata by Bechyné in 1955. We dissected genitalia from four possibly 50–100 year old specimens of D. carinata from central Asia with the following four identification labels: “Diorhabda sulphurea Reitt” (Karatack [Karatag], Tajikistan; DEI), “D. sulfurea Rt” (Tshcardsui [Turkmenabat], Turkmenistan; NMPC), “sulphurea” (Hissar [Hisor], Tajikistan; MSNM), and “Diorhabda sulphureus Reitter det. H. Bollow 1939” (Aulie–Ata [Taraz], Kazakhstan; also with label “D. e. carinata Fald. det. J. Bechyné 1955”; NHMB). The specimen from Karatag, Tajikistan also bore the label “m.n.sp. [manuscript nova species] sulphurea” indicating that “D. sulphurea” was an unpublished manuscript name, probably of the German coleopterist Edmund Reitter who

68 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS actively published ca. 1870–1915. We find no publication of the manuscript name “D. sulphurea”. In any case, Faldermann’s (1837) description of D. carinata precedes Reitter’s works. In the field in central and southwest Asia, it would be useful to distinguish live adults of D. carinata from the sympatric and partially syntopic species D. carinulata and D. meridionalis. Differences in minimum and maximum sizes reached by males and females of D. carinata compared to D. carinulata and D. meridionalis can aid in identification of some specimens (see Table 2). In some living females, the internal sternite VIII appears as a dark Y-shaped area visible through the translucent last visible abdominal sternite (see Lewis et al. 2003b, Fig. 1G). Female D. carinata often have the lobes of internal sternite VIII both pointed and curved inward (Fig. 40—Baghdad, Ashgabat), a condition not seen in D. carinulata and D. meridionalis (Figs. 42–43). As discussed below, a high incidence of Diorhabda egg masses on Tamarix bark (Milbrath et al. 2007) also can indicate the presence of D. carinata. Common Name. The vernacular name “larger tamarisk beetle” refers to the statistically significant larger mean size of D. carinata compared to all other species in the D. elongata group. Among the D. elongata group, only D. carinata exceeds 7 mm in length in males and 8 mm in length in females (Table 2). Diorhabda carinata is especially larger than the two species with which it is moderately to partially sympatric and syntopic, D. carinulata and D. meridionalis. Biology. Host Plants. Because of the misapplication of the name D. elongata to both D. carinata and D. carinulata over a wide area of west and central Asia, we cannot conclusively distinguish biological literature that refers to D. carinata alone. Diorhabda carinata is more commonly collected than D. carinulata in the following areas where we consider field observations to primarily involve D. carinata (see Unconfirmed Records above): eastern Georgia (Lozovoi 1961), Azerbaijan (Samedov and Mirzoeva 1985), southern Turkmenistan (Myartseva 1995, 1999, 2001), and Tajikistan (Kulinich 1962). All previous reports of hosts for D. carinata were made under the name D. elongata. D. carinata (as D. elongata) feeds upon Tamarix ramosissima and T. smyrnensis (as T. hohenackeri Bunge) in eastern Georgia (Lozovoi 1961), T. meyeri Boissier and T. smyrnensis in Azerbaijan (Samedov and Mirzoeva 1985), and T. ramosissima, T. hispida Willdenow and T. arceuthoides Bunge in Tajikistan (Kulinich 1962) (see Unconfirmed Records above) (Table 1). We also examined specimens of D. carinata collected from T. ramosissima near Shelek (Chilik), Kazakhstan by I.D. Mityaev and R. Jashenko, and in Chuli Canyon, Turkmenistan by C.J. DeLoach. Tamarix sp. is recorded as a host for material we examined of D. carinata collected from Shorkot, Pakistan. We report the following four new host records from our dissected material: T. aralensis Bunge from Dry Sport Lake, Ashgabat, Turkmenistan collected by S. Myartseva; T. aucheriana (Decaisne) Baum from Village Danata, Turkmenistan collected by C.J. DeLoach; T. aphylla from Kund, Pakistan (as D. elongata in Habib and Hasan [1982]); and T. ramosissima/T. chinensis from near Borger and Seymour, Texas. We also examined an adult collected on T. cf. indica Willdenow (as T. cf. troupii Hole) from Nomal, Pakistan. Dalin et al. (in press) found that D. carinata (as D. e. carinata) from Uzbekistan preferred T. parviflora to a similar degree as T. ramosissima in multiple-choice field cage studies in southern California, making T. parviflora a potential new host. Diorhabda carinata shares four of its eight known Tamarix spp. hosts with D. carinulata: T. ramosissima, T. arceuthoides, T. hispida, and T. aralensis (Table 1). Diorhabda carinata was collected syntopically from the same trees with D. carinulata on T. aralensis at Dry Sport Lake (nr Ashgabat), Turkmenistan, and T. ramosissima near Shelek, Kazakhstan and Quaragum Canal (nr Ashgabat), Turkmenistan. In no-choice larval host suitability studies, D. carinata larvae from Uzbekistan can only survive to adulthood on plants of the order Tamaricales, including Tamarix (Tamaricaceae) and, to a generally lesser degree, on three North American Frankenia spp. (Frankeniaceae): F. s al in a , F. johnstonii, and F. jamesii (Milbrath and DeLoach 2006a, Herr et al. in prep.). Multiple-choice adult oviposition studies in field cages (Milbrath and DeLoach 2006a) reveal that the three North American Frankenia spp. provide little attraction for oviposition by D. carinata compared to Tamarix. In field cage no-choice studies, oviposition by D. carinata on F. jamesii and F. johnstonii was not different from non-host coyote willow (Salix exigua) and adults experienced increased mortality compared to T. ramosissima × T. chinensis treatments (Milbrath and

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 69 DeLoach 2006a). Among invasive North American Tamarix spp. in large field cages, adult D. carinata oviposited as much on T. aphylla as on T. ramosissima, T. parviflora or T. chinensis × T. canariensis/T. gallica, and oviposited significantly more on T. aphylla than on T. canariensis/T. gallica. However, in a later similar multiple-choice field cage test among Tamarix spp., Milbrath and DeLoach (2006b) found that D. carinata oviposited significantly less on T. aphylla than on all other Tamarix, including T. ramosissima × T. chinensis, T. ramosissima × T. canariensis/T. gallica, T. canariensis/T. gallica and T. parviflora. In less discriminating no-choice field cage studies, D. carinata accepted T. aphylla for oviposition to the same degree they accepted T. ramosissima × T. chinensis (Milbrath and DeLoach 2006b). Tamarix aphylla is at moderate risk of damage by D. carinata in the field and it is difficult to predict to what degree D. carinata would damage T. aphylla, especially in the absence of other Tamarix spp. (Milbrath and DeLoach 2006b). Frankenia is at low risk to damage from D. carinata (Milbrath and DeLoach 2006a). Risk of damage to both T. aphylla and Frankenia by D. carinata is probably much lower when these plants are not in the proximity of preferred Tamarix spp. (e.g., Blossey et al. 2001). Ecology and Phenology. Kulinich (1962) found D. carinata (as D. elongata) damaging leaves and young shoots of tamarisk in Tajikistan, where it occurs from May to September. According to our data from examined material, D. carinata was collected as early as 23 April in Tajikistan. In southern Tajikistan, D. carinata (as D. elongata) can have four generations (Pripisnova 1965). Samedov and Mirzoeva (1985) report D. carinata (as D. elongata) as “frequently badly damaging” bushes of both T. meyeri and T. smyrnensis in Azerbaijan, where it has three generations from April to October. Lozovoi (1961) commonly found D. carinata (as D. elongata) on tamarisk throughout eastern Georgia from 1959–1960, but never in sufficient quantity to damage the plants. We have seen collection records from 18 April to 3 August in Georgia. Diorhabda carinata severely defoliated tamarisk at Dry Sport Lake, near Ashgabat, Turkmenistan in August of 1998, and this was photographed by A. Knutson (for photo, see DeLoach et al. 2003b, Fig. 3) (see Material examined, specimen no. GSWRL 2005-25). We also found D. carinulata at this site in 1997 and 1998, but it was in much lower numbers than D. carinata. Our collaborator, Myartseva (1999) made season long observations of D. carinata (as D. elongata) at Dry Sport Lake in 1999. Five generations were observed, with the overwintered adults emerging and copulating in mid-March, the first eggs appearing in April, and the fifth generation adults emerging in late August and going into diapause. In July and August, temperatures often reached ca. 40°C during many days in which adults and larvae were absent, and this absence may be associated with a period of possible aestivation in the pupal or adult stage. Collection dates from our examined material range from 1 April to 12 September in Turkmenistan, 7 April to 26 September in Uzbekistan, and 9 April to 10 October in Iraq and Iran. Milbrath et al. (2007) found that adult D. carinata (as D. elongata from Qarshi, Uzbekistan) overwintering at Temple, Texas had ca. 60–80% survival from early November through the beginning of March but survival dropped precipitously to ca. 10% by mid-March when the tamarisk leaves were just budding. Overwintered adults began ovipositing in late March giving rise to five generations and a partial sixth generation. Fifth generation adults emerging in early September oviposited for several weeks before ceasing oviposition in November when they appeared to enter diapause. Five generations were also found in the field at Lake Kemp near Seymour, Texas in 2008 where adults were released in late April and the fifth generation adults emerged in the field by late September. Hundreds of adults could be easily found feeding in this area through 9 October. In late July, a group of ca. 2,000 adults emerged and congregated on a group of large tamarisks and over the course of 12 days appeared to migrate en masse east northeast for 46 meters, congregating on certain tamarisk trees, first at 18 meters and depositing eggs and then moving another 46 meters. (C. Randal, pers. comm.). Milbrath et al. (2007) observed that D. carinata may deposit as much as one-third of its egg masses on the bark of trunks and branches of tamarisk. In contrast, D. carinulata, D. sublineata, and D. elongata, all deposit their eggs on tamarisk leaves, with the exception of D. sublineata rarely ovipositing on tamarisk stems (Milbrath et al. 2007). A high incidence of Diorhabda eggs found on the bark of trunks and branches of tamarisk can indicate the presence of D. carinata in areas of sympatry with other tamarisk beetles.

70 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Development and Reproduction. Milbrath et al. (2007) found that, at 28°C, D. carinata (as D. elongata from Qarshi, Uzbekistan) had a development time of 18.6 days from egg to adult (with 73% survival), a fecundity of 233 eggs, and a population doubling time of 5.7 days. These values were all very similar to those found for D. carinulata (Turpan and Fukang), D. elongata (Crete), and D. sublineata (Tunisia) (all as D. elongata) in the same study. Natural Enemies. The tachinid Erynniopsis antennata (Rondani) is reported as a parasite of larvae and pupae of D. elongata in Ashgabat, Turkmenistan (Richter and Myartseva 1996). Diorhabda elongata does not occur in Turkmenistan and this record probably should refer to D. carinata which is generally much more abundant than D. carinulata in Ashgabat. An adult parasitoid wasp, Baryscapus diorhabdivorus Gates & Myartseva (Hymenoptera: Eulophidae), was found inside an adult D. carinata collected from Ashgabat, Turkmenistan in late September. Previous reports of adult B. diorhabdivorus from both larvae and adults of D. elongata in Ashgabat (Gates et al. 2005) should also refer to D. carinata. The ground beetle, Lebia holomera Chaudior (Coleoptera: Carabidae), preys especially upon leaf beetles of the genus Diorhabda in south-central Asia and in the eastern Caucasus (Kyrzhanovskiy 1965), areas where D. carinata is common. The protozoan mircosporidian parasite, Nosema sp., was found in adult D. carinata originating from two collections at Ashgabat, Turkmenistan (shipments GSWRL(CJD)1996-28 [identified by T. Poprawski] and GSWRL(CJD)- 1997-12 [identified by G.M. Thomas]). The fungal pathogen B. bassiana was also found in adult D. carinata from Ashgabat, Turkmenistan (shipment GSWRL(CJD)1996-28) (identified by T. Poprawski). Biogeography. Comparative. Diorhabda carinata differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) primarily continental with distribution usually greater than 500 km from the oceans and ranging to 2,900 m elevation; (2) usually found in warm temperate desert and grassland biomes; and (3) latitudinal range of 30–44°N and most common from 36–41°N (Table 7, Figs. 51–52). Diorhabda carinata is moderately sympatric with D. carinulata, their ranges overlapping in a large area that includes Azerbaijan and central Asia (northern Iran, Turkmenistan, Uzbekistan, Kazakhstan, Tajikistan, Kyrgyzstan, and western China), where both species have been referred to as D. elongata (see synonymy above; e.g., Ogloblin 1936, Yakhontov and Davletshina 1955, Kulinich 1962, Sinadsky 1968, Lopatin 1977a, Samedov and Mirzoeva 1985, Lopatin et al. 2004) (Map 1; Table 8). Diorhabda carinulata differs from D. carinata in ranging further north to 49°N and in being more common from 42–44°N. Conversely, D. carinata is more commonly collected than D. carinulata in some more southern areas from 35–42°N, especially in the deserts of the Transcaucasus and grasslands of Tajikistan. However, D. carinulata appears to be more common than D. carinata from 31–34°N in deserts of eastern Iran (Map 6). Further studies are needed to better understand the differing climatic preferences of these species. Diorhabda carinata and D. carinulata are also syntopic in some areas, having been collected together in the same series from the same tamarisk trees at three localities: Dry Sport Lake and Quaragum Canal, near Ashgabat, Turkmenistan, and Shelek, Kazakhstan. Diorhabda carinata and D. carinulata were also collected together in the same series (and probably also the same tamarisk trees) at Ordubad, Azerbaijan, and three location in Uzbekistan: Buxoro (5 km northwest), Qarshi (10 km west), and 140 km south of Toshkent. These species were collected in the same locations but differing series at Qobustan, Azerbaijan; Golestan Biosphere Reserve, Iran; Sarytogay Forest, Kazakhstan; and Yining, China. Care must be taken to separate D. carinata and D. carinulata in laboratory colonies begun from areas of sympatry and syntopy in Central Asia. Diorhabda carinata is partially sympatric with D. meridionalis in western Iran and Syria (Maps 1, 6; Table 8). These species are probably also syntopic over some areas, having been collected together in the same series at Halab, Syria and at four locations in western Iran: Jolow Gir, 34 km SE of Omidiyeh, Shushtar, and 30 km NNE of Borazjan. Diorhabda meridionalis differs from D. carinata in being maritime and common further south at 26–31°N (Figs. 51–52). In the small sample sizes in which D. carinata and D. meridionalis were collected together from 29.5–36°N, D. carinata was a little more abundant than D. meridionalis north of 30°N while D. meridionalis was a little more abundant south of 30°N (at 30 km NNE Borazjan). Diorhabda carinata is most similar to D. meridionalis and D. carinulata in terms of inhabited biomes (Fig. 53).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 71 Diorhabda carinata is marginally sympatric with D. sublineata, their distributions meeting in Baghdad, Iraq. Diorhabda sublineata differs from D. carinata in being primarily maritime with a strong presence in the Mediterranean biome, and in ranging further south to 16°N where it is most common from 31–35°N. Diorhabda carinata is most biomically different from D. sublineata, the species with which it is most morphologically similar (Tables 9 and 10; Fig. 53). Diorhabda carinata is probably marginally sympatric with D. elongata in eastern Turkey, western Syria, and, possibly, southern Russia, Georgia and Azerbaijan (Map 1). Diorhabda elongata differs from D. carinata in being maritime with a strong presence in the Mediterranean biome (Figs. 51–52). Descriptive. Diorhabda carinata is primarily found in the south central Palearctic realm, but two collection locations in northern Pakistan (Shorkot and Islamabad) fall within the borders of the Indo-Malayan realm (Maps 1, 3, 6). Most collections originate between 37° and 41°N in two biomes of Central Asia: the Deserts and Xeric Shrublands (ca. 0–800 m elevation) and Temperate Grasslands, Savannas and Shrublands (ca. 250–1,600 m) (Map 3, Table 9). Reports of D. carinata damaging tamarisk (Kulinich 1962, Samedov and Mirzoeva 1985, Myartseva 1999) originate from these two biomes in this region. Primary ecoregions for D. carinata in the Temperate Grasslands, Savannas and Shrublands biome from 37–41°N include the Gissaro–Alai Open Woodlands (ca. 250–450 m) and the Alai–West Tian Shian Steppe (ca. 400–1,600 m) in south central Kazakhstan and western Uzbekistan, Tajikistan and Kyrgyzstan (Map 3). Primary ecoregions with D. carinata in the Deserts and Xeric Shrublands biome from 37–41°N include the Azerbaijan Shrub Desert and Steppe (ca. 0–400 m) and the Central Asian Southern Desert in south central Kazakhstan, southern Uzbekistan and Turkmenistan (ca. 200–350 m). Diorhabda carinata occurs more frequently in the Temperate Conifer Forests biome than any other Diorhabda species, where it may be found from 33–41°N in northwestern Iran, eastern Turkey and northern Pakistan (ca. 550–1,600 m; Maps 3 and 6; Table 9). Other biomes with D. carinata from 37–41°N include the Temperate Broadleaf and Mixed Forests of northern Iran and northeast Turkey (ca. 350–550 m) and Montane Grasslands and Shrublands (Kopet Dag Woodlands and Forest Steppe ecoregion) along the border of Iran and Turkmenistan (ca. 200–850 m). The westward distribution limit of D. carinata in Turkey and Ukraine corresponds well with the westward distribution limit of one of its host plants, T. ramosissima (Map 3). The distribution of the host T. meyeri, which ranges across deserts from Azerbaijan eastwards to Uzbekistan and Afghanistan (Rusanov 1949, Baum 1978; not shown), corresponds to the center of distribution of D. carinata in central Asia. The distribution of the host plant T. aralensis approximately coincides with the distribution of D. carinata in deserts of its southern range in Iran and Iraq as well as over its north-central range in Turkmenistan, Uzbekistan and Tajikistan (Map 3). The northern distribution of the host T. aucheriana also coincides with the range of D. carinata in Iraq and Iran (Map 3). The north central distribution of the host T. arceuthoides (Rusanov 1949, Baum 1978, Browicz 1991; not shown) coincides with the heavy areas of occurrence of D. carinata in its northeastern range in Tajikistan and southern Uzbekistan. Tamarix arceuthoides is a very common Tamarix from eastern Iraq to Tajikistan (100–3,000 m elevation) that occurs more commonly in mountainous rocky or pebbly substrates than does T. ramosissima (Rusanov 1949, Liu 1987, Browicz 1991) and prefers lower salinity habitats compared to T. aucheriana (Leonard 1992). We find no reports of damage by D. carinata from the southern portion of its range, at 29–37°N (Map 6). Habib and Hasan (1982) studied Tamarix insect herbivores in northern Pakistan, but did not report any damage by D. carinata (as D. elongata) from 33–36°N. South of 37°N, D. carinata is primarily found in the Deserts and Xeric Shrublands biome of Iraq, Afghanistan, and Pakistan (ca. 50–1,500 m), but it also occurs in the Temperate Broadleaf and Mixed Forests and Temperate Conifer Forests biomes of western and northern Iran (ca. 1,150–1,900 m) (Maps 3 and 6; Table 9). Potential in Tamarisk Biological Control. Summary. The larger tamarisk beetle is apparently establishing near Seymour, Texas (Map 7). Diorhabda carinata may be the most effective tamarisk beetle for control of T. ramosissima/T. chinensis in the temperate grasslands biome, including the Western Short Grasslands and Central and Southern Mixed Grasslands, and temperate conifer forest biome which includes the Arizona Mountains Forests of New Mexico and Arizona (Map 13). Diorhabda carinata may co-dominate with D.

72 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS carinulata in some temperate warm desert areas such as the Trans-Pecos Chihuahuan and Mojave deserts and southern portions of the Great Basin Shrub Steppe and Colorado Plateau shrublands (Map 13). Discussion. The larger tamarisk beetle damages Tamarix in west to central Asia from latitude ca. 38–41°N and ca. -10–1,000 m elevation in Azerbaijan (Samedov and Mirzoeva 1985), Turkmenistan (Myartseva 1999), and Tajikistan (Kulinich 1962). Diorhabda carinata attacks T. ramosissima throughout the northern portion of its range in Georgia (Lozovoi 1961), Turkmenistan (present study), Tajikistan (Kulinich 1962) and Kazakhstan (present study), and damages T. smyrnensis, a close relative of T. ramosissima, in Azerbaijan (Samedov and Mirzoeva 1985). We expect D. carinata would attack and probably damage and control T. ramosissima/T. chinensis in North America. It has a moderate risk of damaging T. aphylla (Milbrath and DeLoach 2006b) and very low risk of damaging Frankenia (Milbrath and DeLoach 2006a), and both these risks are probably much reduced at less proximity to preferred Tamarix spp. (e.g., Blossey et al. 2001). Diorhabda carinata appears to be establishing on Lake Kemp, near Seymour, Texas where it was released in April, 2008 and defoliated more than 0.2 ha by August and was common over a 0.8 ha area (C. Randal, pers. comm.; Maps 7 and 13). Diorhabda carinata defoliated portions of a few saltcedar trees in the summer of 2008 following its spring release at Matador Wildlife Management Area (WMA) near Paducah, Texas (Mike Janis, Texas Parks and Wildlife Department, Matador WMA, pers. comm.). It appeared to be weakly established on the Canadian River near Borger, Texas where it was first released (under permit as D. elongata) in July 2006, but populations could not be found in 2008 (J. Michels and E. Jones, pers. comm.). D. carinata was also released in west Texas at Roaring Springs, Rotan, and Guthrie in the summer of 2008, but it has not yet established at these sites (A. Knutson, pers. comm.). D. carinata was caged in the Mojave Desert at Camp Cady, California in 2008 and permissions are being sought for release at this site in 2009 (T. Dudley, University of California, Santa Barbara, CA, pers. com.). Northern climatypes of larger tamarisk beetles originating from the Desert and Xeric Shrublands and Temperate Grasslands and Shrublands biomes between 37–41°N probably have the greatest potential to damage tamarisk in corresponding latitudes and biomes within 0–1,000 m elevation in North America (Map 13). Areas of primary ecoregions matching these criteria are the Western Short Grasslands in western Kansas, portions of the Great Basin Shrub Steppe and Colorado Plateau shrublands, and some of the extreme northern Mojave Desert (Map 13). Elevations above 1,100 m from 37–41°N should be suitable, but in some cases suboptimal, for D. carinata. Milbrath et al. (2007) found that a northern climatype of D. carinata from 38°N in Uzbekistan suffered higher overwintering mortality in early March at 31°N in Temple, Texas than did D. sublineata and D. elongata. Higher mortality in the northern D. carinata climatype may have been related to asynchronization in the breaking of adult quiescence (not diapause, which probably ends in early winter in all the tamarisk beetles) with bud break in tamarisk as signaled by late winter/early spring temperatures. At 31°N in Temple, higher temperatures in late winter/early spring may occur several weeks before tamarisk bud break (temperatures begin to gradually warm before bud break), but at 38°N these same higher temperatures may occur later in the season and be more closely associated with tamarisk bud break (temperatures may more sharply warm up before bud break). A more southern climatype of D. carinata (see below) may be better synchronized in responding to higher temperatures as a signal to ending of winter quiescence at 31°N. Several North American ecoregions correspond to the most northern areas inhabited by D. carinata in its native distribution from 41–45°N (Map 13). These regions include northern portions of the Western Short Grasslands and southern portions of the Wyoming Basin Shrub Steppe (Map 13). Much of these northern areas may be more suitable for D. carinulata. Southern climatypes of D. carinata (Map 6) may be best suited to ecoregions in the Temperate Grasslands, Savannas and Shrublands biome between 31–37°N (Map 13). These include the southern portions of the Western Short Grasslands in eastern New Mexico and northwest Texas and southwest portions of the Central and Southern Mixed Grasslands in north central and western Texas (Map 13). Several temperate warm desert regions in North America correspond to native habitats for southern D. carinata climatypes in the Desert and Xeric Shrublands biome between 31–37°N. These desert ecoregions include southern portions of the Colorado Plateau Shrublands, portions of the Mojave Desert in southern California and Nevada, and the

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 73 Trans-Pecos Chihuahuan Desert in Texas, New Mexico and northern Chihuahua, Mexico (Map 13). Larger tamarisk beetles might damage tamarisk in some of these southern grassland and desert ecoregions. Diorhabda carinata appears to be the best suited species to the Temperate Conifer Forests among the D. elongata group (Tables 9 and 13). This would make D. carinata the best suited species to tamarisk invaded riparian areas of the Arizona Mountains Forests ecoregion of Arizona and New Mexico, such as along the Rio Hondo within the pinyon-juniper woodlands near Hondo, New Mexico (Map 13). Southern maritime subtropical desert areas of 31–29°N, such as the Sonoran Desert, may not be as suitable for D. carinata as for other more southern sibling species (Map 13, Table 7).

Diorhabda sublineata (Lucas, 1849) REVISED STATUS subtropical tamarisk beetle (Figs. 5, 6, 16, 21, 26, 31, 36, 41, 45)

Galeruca sublineata Lucas, 1849:542 (Type locality: Hippône [Annaba], Algeria; as Galleruca). Galeruca elongata: Reiche and Saulcy, 1858:42 (part; France, Egypt, Algeria; as Galleruca); Joannis, 1866:83 (part, monograph, France, as Galleruca). Diorhabda elongata: Heyden et al., 1891:375 (part, catalog for Europe and Caucasus, southern Europe), Bedel, 1892:158 (part, France, as Dirrhabda); Weise, 1893:635 (part, Algeria), 1925:225 (Egypt); Corréa de Barros, 1924:9 (part, Portugal); Peyerimhoff, 1926:359 (part, Algeria, Senegal, hosts); Laboissière, 1934:54 (part, France, Senegal, host); Ogloblin, 1936:79 (part; European and African coast of Mediterranean Sea, south to Senegal); Normand, 1937:126 (Tunisia); Kocher, 1958:109 (Morocco, to 2,000 m); Hopkins and Carruth, 1954:1129 (part, Spain, host); Jolivet, 1967:331 (part, Morocco, Mediterranean, hosts); Torres Sala, 1962:327 (part, Comunidad Valenciana, Spain); Petitpierre, 1988:93 (part, Spain); Gruev and Tomov, 1998:70 (part, Mediterranean); Kovalev, 1995:78 (part, southwest Palaearctic); Campobasso et al., 1999:145 (part, host, Europe and Middle East); Anonymous, 2001:52N (part, Africa); Chatenet, 2002:223 (part, France, Spain, Morocco, Algeria, Tunisia); DeLoach et al., 2003a:230 (part, Tunisia [Sfax]), 2003b:126 (part, host range, North Africa, France), (2008, in prep.) (part, Sfax, Tunisia); Milbrath et al., 2003:225 (part, Sfax, Tunisia); Riley et al., 2003:69,189 (part, catalog of North America [introduced]); Warcha- lowski, 2003:328 (part, taxonomic keys, Mediterranean Region); DeLoach and Carruthers, 2004a:13, 2004b:311 (part, Tunisia [Sfax]); Lopatin et al., 2004:127 (part, Mediterranean); Dudley, 2005a:13, 2005b:42N (part, biological control, ex: Tunisia); Milbrath and DeLoach, 2006a:32, 2006b:1379 (part, host specificity, Sfax, Tunisia); Dudley et al., 2006:137 (part, host range, Sfax, Tunisia); Milbrath et al. (2007). (part, biology, Sfax, Tunisia); Bean and Keller (in prep.) (part, diapause induction, Sfax, Tunisia); DeLoach (2008); Moran et al. in press (host range of D. sublin- eata [ex: Sfax, Tunisia] × D. elongata [ex: Crete] hybrid); Thompson et al. (in prep.) (part, laboratory hybridization, Sfax, Tunisia). Diorhabda elongata var. sublineata: Weise, 1893:1132 (part, Algeria). Diorhabda sublineata: Boehm, 1908:68 (Egypt, as Dirrhabda). Diorhabda elongata ab. sublineata: Weise, 1924:78 (part, world catalog, Algeria); Winkler, 1924–1932:1307 (part, Palearctic catalog, Algeria); Laboissière, 1934:53 (part, France, Algeria); Normand, 1937:126 (part, Tunisia); Alfi- eri, 1976:233 (Egypt, on Tamarix spp.); Warchalowski, 2003:328 (part, taxonomic keys, Mediterranean Region). Diorhabda elongata ab. carinata: Laboissière, 1934:54 (part; Narbonne, France). Diorhabda elongata ab. bipustulata Normand, 1937:126 (Type locality: , Tunisia) (NEW SYNONYM); Wil- cox, 1971:63 (world catalog). Diorhabda elongata sublineata: Gressitt and Kimoto, 1963a:407 (part; North Africa, West Asia; not Asia Minor, China, and Mongolia); Wilcox, 1971:63 (part, world catalog, North Africa, West Asia); DeLoach et al., 2003b:126 (part, host range, North Africa).

Male. Genitalia. Male D. sublineata may be distinguished from all other species in the D. elongata group by the unique presence of a connecting endophallic sclerite (CES) from the palmate endophallic sclerite (PES) to the elongate endophallic sclerites (EES) (Figs. 16, 21, 26, 31). In weakly sclerotized teneral adult D. sublineata, the CES can appear to evanesce and be faint. Diorhabda elongata, D. carinata, D. carinulata and D. meridionalis all lack the CES (Figs. 14–15, 17–20, 22–25, 27–30, 32–33), but darkly sclerotized D. carinata may uncommonly bear a faint lateral line in place of a CES. Additional characters of the EES and PES, similar to those found in D. carinata, can be used to separate D. sublineata from D. elongata, D.

74 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS carinulata, and D. meridionalis. The PES of D. sublineata (Figs. 16, 31) always bears a strong lateral appendage and the distal margin is truncate-serrate with two to six (commonly three to four) usually distal spines, a maximum of one spine being subdistal. In contrast, the PES of D. carinulata (Figs. 17, 32) and D. meridionalis (Figs. 18, 33) lacks a lateral appendage (but it may bear a lateral notch) and the distal margins of the PES are narrowly rounded and generally smooth with one or two small subdistal spines that sometimes project beyond the distal margin. The PES of D. elongata also lacks a lateral appendage and is usually rounded with one to five usually subdistal spines with a maximum of two of these spines being distal (Figs. 14, 29). The elongate endophallic sclerite in D. sublineata always bears a basally pointed lateral appendage (LA) or lateral notch (LN) serving as a point of attachment for the CES (Figs. 16, 21), but these are always lacking in D. elongata, D. carinulata, and D. meridionalis (Figs. 14, 17–19, 22–23). The EES of D. carinata only occasionally bears a lateral appendage (Figs. 20—Artvin) or a lateral notch (Fig. 20—Ashgabat). In D. sublineata, the spined area of the EES extends greater than or equal to 0.31 times (or greater than about one third) the length of the EES (Figs. 16, 21, 48; SL). In contrast, the spined area of the EES is confined to less than or equal to 0.16 times (or less than about one fifth) the length of the sclerite in D. elongata (Table 3; Figs. 14, 19, 24, 48). In D. sublineata, spines of the EES are often irregularly spaced along the blade with conspicuous gaps (Fig. 21). In D. carinulata and D. meridionalis, spines along the EES are usually evenly and closely spaced along the blade (Figs. 22–23). The EES of D. meridionalis additionally bears a hooked apex that is absent in D. sublineata. Measurements. See Tables 2 and 3. Female. Genitalia. Female D. sublineata may be distinguished from all other members of the D. elongata group except D. carinata by their triangulate vaginal palpi (VP) that are wider than long with a width to length ratio (LP/WP) of 0.46–0.85 (n = 19) (Fig. 36; Table 4). In contrast, the vaginal palpi are broadly rounded with a length to width ratio of 0.94–1.36 in the allopatric D. carinulata (Fig. 37) and D. meridionalis (Fig. 38; Table 4). The vaginal palpi are also broadly rounded in D. elongata (Fig. 34). In addition, the width of the widest lobe of the stalk (WLS) of internal sternite VIII (IS VIII) is often larger in D. sublineata (range 0.08–0.18 mm; Fig. 36) compared to D. elongata (range 0.06–0.11 mm; Fig. 34). Some female D. sublineata can be distinguished from D. carinata by having the tips of the lobes (TL) of the stalk of IS VIII either not curved or curved outward toward the apical lobe and the tips either rounded or quadrate (Fig. 41 – Ndiol, Perpignan, Kom Ombo), a state never found in D. carinata. Some D. carinata can be distinguished from D. sublineata by having the tips of both lobes of the stalk of IS VIII strongly curved inward at the tip and the tips either pointed or rounded (Fig. 40 – Baghdad, Ashgabat, and Pul-e Charki). Female D. sublineata and D. carinata with at least one lobe of the stalk of IS VIII curved only slightly inward (Figs. 35–36, 40—Lagodekhi and Ardanuc, 41—Biskra and Tamri) are indeterminable to species, except on the basis of geographic distribution outside of the area of known sympatry in Iraq. Measurements. See Tables 2 and 4. Coloration. In D. sublineata, subsutural and submarginal elytral vittae are often present, extending well into the basal half of the elytra (Fig. 5) and live specimens are tannish-yellow in hue, lacking any greenish- yellow tinting (Fig. 6). This is in contrast to D. elongata in which the elytral vittae, if present, are confined to the apical half of the elytra (Fig. 1) and in which live specimens possess an olivaceous hue from greenish- yellow tinting (Fig. 2). Diorhabda carinata differs from D. sublineata in more often lacking elytral vittae (Fig. 3) and in having some degree of greenish-yellow tinting in veins of the elytra (Fig. 4). Living D. sublineata (Fig. 6) and D. carinulata (Fig. 8) are very similar in coloration, and Chen (1961) notes the similar appearance of dead D. carinulata (as D. e. deserticola) and D. sublineata (as D. e. ab. sublineata). Type material. Specimens from Lucas’s collections, which should include the type specimen(s) for D. sublineata, should be found at MNHN (Groll 2006). The curator at MNHN communicated intent to inform us of the status of the type material and perhaps lend it for examination, but after four years, we have not heard of the status of the Lucas type material. Once it can be ascertained that the type material is lost, a neotype should be designated using a dissected male specimen from near the type locality of Annaba, Algeria. We studied the original description by Lucas (1849), possibly based on several individuals he stated were collected at

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 75 Annaba, with his accompanying color habitus illustration. We also studied specimens from the broad vicinity of the type locality in Algeria and Tunisia. The location of type specimens for D. e. ab. bipustulata Normand (1937) is unknown and we studied the original description and specimens from the vicinity of the type locality of Kairouan, Tunisia. Material examined. 89♂♂ dissected (diss.), 51♀♀ diss., 170♂♂, 191♀♀, 9 unsexed specimens. ALGERIA: 2♀♀ diss., [specific location not given], Reitter, NMPC [2004-21, 25]; 1♂, [specific location not given], Uyttenboogaart, Coll. Dr. D.L. Uyttenboogaart, ZMAN; 1♂ diss., 2♂♂, 3♀♀, Abadla [31.0167°N, - 2.7333°W], Kouril, P5/46/62, 4358a, NMPC [2004-03]; 1♂ diss., 6♀♀, Abadla, V. Zoufal, Kouril, P5/46/62, 4358a, D. elongata a. sublineata [ZMHB], NMPC [2004-14], ZMHB [1♀]; 1♀, Abadla, ZMHB; 1♂ diss., 7♂♂, 8♀♀, Alger [36.76305°N, 3.05055°E], environs, Ch. Lallemam, Diorhabda elongata var. carinata det V. Laboissière 1939 [8♂♂, 8♀♀], D. e. var. sublineata det V. Laboissière 1939 [1♀], IRSNB [2006-02]; 1♂, Algir [Alger], coll[ection] Reitter, D. elongata, HNHM; 1♂ diss., Argel [Alger], St. Perez Arcas, Galeruca elongata, MNMS [2003-01]; 1♂ diss., Biscra [Biskra; 34.85°N, 5.73333°E], Diorhabda elongata det I.K. Lopatin, Coll. Lichtnekert, HNHM [2003-03]; 1♂ diss., 1♂, 2♀♀, Biskra, environs, Obenb., NMPC [2004- 22]; 1♂ diss., 1♀ diss., 2♂♂, 6♀♀, Biskra, C.J. Dixon, Museum Lieden, RBCN [2003-08, 21]; 1♂ diss., 2♀♀ diss., 2♂♂, 1♀, Biskra, G.C. Champion, G.C. Champion Coll. B.M. 1927–409, BMNH [2003-03, 2003-11, 2005-01]; 1♀ diss., 1♀, Biskra, 1-III-1931, Dr. R. Meyer, NHMB [2005-08]; 2♀♀ diss., 3♀♀, Biskra, 30-III- 1985, M. Bergeal, MBPF [2003-01, 02]; 1♂, 3♀♀, Biskra, P. Jolivet, IRSNB; 1♀, Biskra, J. Sahlb., MZHF; 1♂ diss., Blidah [Blida; 36.41667°N, 2.828889°E], V-1867, H. Clark, 65-56, BMNH [2003-08]; 1♀ diss., Boghari [Boghni; 36.5422°N, 3.9531°E], V-1897, Dr. A. Chobaut, NMPC [2004-01]; 1♀ diss., Qued Hamman [Oued Hammam (Mascara State)], 10 km south Perregaux [Mohammadia] [35.50806°N, 0.040833°E], 9-V-1964, Eckerlein, Coll[ection] K.H. Mohr, DEI [2005-05]; 1♂ diss., 8♂♂, 13♀♀, Teniet el Had [Theniet el Had] to Affreville [Khemis Miliana] [app. location; 36.13950°N, 2.20749°E], Dr. Vauloger, D. elongata det. Le Moult, IRSNB [2006-07]; EGYPT: 5♂♂, 1♀, [specific locality not given], Reitter, D. sublineata det. V. Laboissière 1913 [HNHM], NMPC [4♂♂, 1♀], HNHM [1♂]; 1♂, [specific locality not given], col[lection] Mus Murray, Fry coll[ection] 1905.100, Galleruca costipennis Dej cat [Dejean catalogue; nomen nudum; Dejean 1837, p. 377], BMNH; 2♂♂, [specific locality not given], ZIN; 1♂ diss., Cairo [Al Qahirah; 29.98333°N, 31.13333°E], Dachor, 29-I-1933, Schatzm. Koch., [D. e. ab.] sublineata det. Burlini 1966, MSNM [2003-17]; 1♂ diss., 1♂, 1♀, Cairo [Al Qahirah], Pyramidi, 3-X-1933, W. Wittmer, D. e. ab. sublineata det. Burlini 1966 MSNM [2003-02]; 10♂♂ diss., 3♀♀ diss., 43♂♂, 50♀♀, Com Ombo [Kom Ombo; 24.45°N, 32.93333°E], XI-1954, D. e. ab. sublineata det. J. Bechyne 1951, H. Hofbrauer, NHMB [2003-01, 04, 08, 2004-05, 06, 07, 08, 09; 2008-01, 02, 03, 04, 05]; 1♂ diss., 9 specimens, Dakhla [25.50722°N, 28.94722°E] MUT, 10-XII-1977, R.T. Simon Thomas, ZMAN [2008-11]; 1♂ diss., 1♂, 1♀, Dakhla Oasis, 4-III-1995, G. Strauss, RBCN [2003-02]; 1♂ diss., 3♂♂, 1♀, Ezbet El Nakl [Izbat an Nakhl; 30.15°N, 31.31666°E], 22-VIII-1933, W. Wittmer, [D. e. ab.] sublineata det. Burlini 1967, MSNM [2003-10]; 1♀ diss., 2♀♀, Fajum [Al Fayyum; 29.30778°N, 30.84°E], U. Sahlb., [no.] 610, MZHF [2003-01]; 1♀ diss., Hawara [pyramid; 29.27389°N, 30.90139°E], 2-V-1904, D. elongata det. L. Burgeon, D. e. var. sublineata det. V. Laboissière 1939, L. Burgeon Collection, IRSNB [2006-06]; 1♂ diss., 1♀, Heluan [Helwan des Bains; 29.85°N, 31.3333°E], 18-IX-1957, dr. Gozmány, in desert on Tamarix, Exc. Egypt Mus. Nat. Hung., D. e. var. sublineata det K. Lopatin, HNHM [2003-07]; 1♀ diss., Ismailia [30.5833°N, 32.2667°E], II, F. Lotte, USNM [2003-17]; 1♂, 2♀♀, Ismailia, F. Lotte, Février, ZIN; 4♂♂, 5♀♀, Ismailia, NMPC; 1♂ diss., 1♂, 5♀♀, Le Caire [Al Qahirah], NMPC [2004-45]; 1♂ diss., 1♂, Port Said [31.2667°N, 32.3°E], VI (June), F. Lotte, USNM [2003-19]; 1♀ diss., 1♀, Sakkarah, [ruins; 29.8457°N, 31.2104°E], Cairo (Gizeh), 19-II-1933, C. Koch, D. elongata det. Burlini 1966, MSNM [2008-03]; 1♂ diss., Siala [Silah; 29.3561°N, 30.9689°E], 21- VIII-1910, collection L. Burgeon, D. e. var. sublineata det. V. Laboissière 1939, IRSNB [2006-01]; 1♀ diss., Wadi Hoff [Ain Elwan Station; 29.86667°N, 31.31667°E], 20-V-1922, Alfieri, [no.] 1972, NHMB [2004-01]; FRANCE: 1♂ diss., 1♂, 3♀♀, Almanane [beach near Hyères; 43.0790°N, 6.1258°E], D. elongata det. V. Laboissière 1939, IRSNB [2006-03]; 1♂ diss., 11♂♂, 22♀♀, Camargue [region], Gallia, L. Puel, coll[ection] Leonhard [DEI], coll. H.K. Mohr [DEI], D. elongata [DEI], DEI [2005-08; 4♂♂, 2♀♀], IRSNB [collection P.

76 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Jolivet; 7♂♂, 20♀♀]; 10 specimens, Ilis [37.8833°N, 21.3833°E], Olympia, Ellas, 2–3-X-1962, Ent. Exe. Zoöl. Mus., ZMAN; 2♂♂ diss., 2♂♂, 2♀♀, Le Lavandou [43.1322°N, 6.3645°E], Exped. Obenb., NMPC [2004-19, 2008-01]; 2♀♀, Le Lavandou, IV-1910, H. Desbordes, D. elongata det. Le Moult, IRSNB; 1♂ diss., 1♂, Le Barcares [42.78333°N, 3.03333°E], 19-V-1973, G.J. Slob, RBCN [2003-05]; 1♂ diss., Stes. Mairies [Les Saintes Maires, 43.45°N, 4.4333°E], Camargue, 11-13-IX-1951, H.H. Weber, NHMB [2003-19]; 1♀ diss., Les Stes.–Maries [Les Saintes Maires], 14-VII, W. Liebmann, DEI [2005-07]; 1♂ diss., 5♂♂, Marseille [43.3°N, 5.4°E], 16-VIII-1952, Hakan Lindb., MZHF [2004-01]; 1♂ diss., Montpellier [43.6°N, 3.833°E], Jentsson, tamarisci, Janis, DEI [2003-17]; 1♂ diss., 1♂, 1♀, Palavas [Palavas–les–Flots; 43.53333°N, 3.93333°E], [no.] 25-11, MBPF [2003-06]; 1♀ diss., Perpignan [42.68333°N, 2.88333°E], Pyr[ennes Mts.], Or[iental], IV-1953, J.V. Bechyné, NHMB [2003-43]; 1♂ diss., 3♂♂, 10♀♀, Salin–de–Giraud [43.4167°N, 4.7333°E], B du Rhône, Camargue, 12-IX-1970 [RBCN 2003-17; 2♀♀ ZMAN], 14-IX-1970 [ZMAN], C. van Nidek, D. elongata [ZMAN], ZMAN, RBCN [2003-17]; 1♂ diss., Séveillé [geocoordinates not locatable], southern France, coll[ection] Bourgoin, D. elongata det. Le Moult, IRSNB [2006-05]; IRAQ: 1♂ diss., Baghdad [33.33861°N, 44.37722°E], IV-1936, Frey, NHMB [2003-31]; MOROCCO: 1♀ diss., Asni [31.25°N, -7.98333°W], 1,000 m [elev.], High Atlas [Mts.], 9-VII-1993, Stn. Sammlung Dieter Siede, D. elongata det. R. Beenen 1995, RBCN [2003-10]; 2♂♂ diss., Driouch [34.9833°N, -3.3833°W], near Midar, Nador [Prov.], 24-VI-1992, J.M. Vela, D. elongata det. Vela 1992 [1♂ diss.], GSWRL [2003-36, 2005-78]; 1♂ diss., Essaouira environs [31.51305°N, -9.76972°W], Ounara, 3-V-1983, S. Doguet, MBPF [2003-08]; 1♂ diss., Goundafa [Talaat n' Yakoub; 30.9833°N, -8.15°W], 1,200 m [elev.], Haut Atlas [Grand Atlas Mts.], 15-30-IV-1933, Schwingenschus, NHMB [2003-18]; 1♂ diss., 2♀♀, Marrakesch [Marrakech; 31.6333°N, -8.000°W], 15-V-1992, Liebegott, D. elongata det. Eber 1995 [1♂ diss., ab. bipustulata], ZMHB [2006-03]; 1♂ diss., 1♀, Marrekech, 21–23-V-1926, Lindberg, MZHF [2008-02]; 1♂ diss., 6♀♀, 2♂♂, Tacheddirt [31.1667°N, -7.85°W] Schwingenschus, NHMB [2003-02]; 1♀ diss., Tamri [30.6978°N, -9.8253°W], Kuste [coast], 31-VII-1993, Stuben, D. elongata det. R. Beenen 1995, RBCN [2003-19]; 2♂♂ diss., 3♀♀, 1♂, Tanger [; 35.78472°N, -5.812778°W], Rolph, Galeruca elongata, tag 1444, DEI [2003-06, 23]; 1♀ diss., Tanger [Tangier], 25–29-IV-1926, Lindberg, 889, MZHF [2008-01]; 1♂ diss., Tanger [Tangier], 1897, MNMS; 2♂♂ diss., 1♀ diss., 5♂♂, 3♀♀, Tarondant [Taroudannt], 12 km east [30.52268°N, -8.759°W], 11-V-1958, C.J. Davis, on Tamarix sp., #83, 58-9590, 58-9591, 58-9592, USNM [2003-01, 02, 03]; 1♂ diss., 1♂, Taroudant [Taroudannt; 30.4833°N, -8.8667°W], 28-V-1985, G. Sama, MRSN [2003-04]; 1♂ diss., Taroudannt, Sous Valley, 18-IV-1990, Z. Kejval, D. elongata, EGRC [2005-06]; 1♂ diss., 3♀♀, Tiz-n-Test Pass [30.83333°N, -8.33333°W], High Atlas [Mts.] 20-VI-1994, Szailles, RBCN [2003-11]; PORTUGAL: 1♂ diss., Lusitania [Portugal, specific locality not given], Reitter, ZMAN [2008- 16]; SENEGAL: 1♂ diss., specific locality not given, Mion, ZMHB [2006-06]; 2♂♂ diss., 1♀ diss., 8♂♂, 2♀♀, Ndiol [Ndiol Nar; 16.14778°N, -16.31306°W], 25 km northeast St. Louis, 15-VIII-1979, P. Jolivet, on T. senegalensis, D. sp. nr. elongata det. S.L. Shute 1979, BMNH [2003-01, 02, 2005-02]; SPAIN: 1♀ diss., [specific locality not given], Lolobregato, NHMB [2003-05]; 2♂♂, 3♀♀, m[eridionalis; southern; specific locality not given], Minsmer, [18]99, ZMAN; 1♂, 1♀, [specific locality not given], L. Miller, coll. Ed. Everts, ZMAN; 1♂ diss., Almeria [36.8833°N, -2.45°W], Tabernas Desert, 21-VII-1965, La Greca, MRSN [2003- 03]; 2♂♂, Andalucia [Comunidad Autonoma de Andalucia; specific locality not given]; coll[ection] Reitter, D. elongata, HNHM; 1♂, 1♀, Andalusia [Andalucia], Baly coll[ection], Galeruca sublineata, BMNH; 1♂ diss., Barcelona [41.3833°N, 2.1833°E], X-1940, F. Monros, USNM [2003-07]; 3♀♀ diss., Barcelona, Farola de Llobregat, 15-VII-1940, F. Monros Coll., USNM [2005-02, 05, 06]; 1♂ diss., Cabo de Gata [El Cabo de Gata; 36.7833°N, -2.2333°W], 4-VI-1977, Hinterseher, D. elongata det. Erber 1985, D. elongata det. R. Beenen 1996, ZMHB [2008-03]; 1♂ diss., 5♂♂, Cordoba [37.83333°N, -4.7667°W], Col del Sur Perez Arcas, G. elongata, MNMS [2004-03]; 1♀ diss., Galicia [Autonomous Community; specific locality not given; estimate approx. locality in Pontevedra Province, the only province of Galicia where Tamarix is noted (Cirujano 1993): 42.0186°N, -8.87510°E], [D.] elongata, NMPC [2005-31]; 2♀♀ diss., Halizia [Galicia Autonomous Community; specific locality not given], [D.] elongata, NMPC [2004-16, 2005-27]; 1♂ diss., Rambla del Grao, Quadix [37.3°N, -3.1333°W], Granada, F.S. Pinero, JOSV68935, GSWRL [2005-77]; 1♂

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 77 diss., 3♂♂, Lorca [37.6667°N, -1.7°W], Murcia, VIII-1943, G. Menor, MNMS [2004-02]; 1♂ diss., Madrid [40.4°N, -3.6833°W], Montarco, Peris Torres, MNMS [2004-01]; 1♀ diss., 14♂♂, 14♀♀, Palamos [41.85°N, 3.13333°E], Catalonien, ZHMB [2003-06]; 1♂ diss., 4♂♂, 3♀♀, Playa de Aro [41.8167°N, 3.0667°E], 2-17- VII-1963, D. elongata det. L. Borowiec, MZLU [2005-04]; 1♀ diss., Sierra de Javalambre [Mt.; 40.1°N, - 1.0°W], Teruel, 1,600m [elev.], 25-VII-1965, La Greca, D. e. subsp. sublineata det. Daccordi 1971, MSNM [2003-04]; 1♂ diss., 2♂♂, 4♀♀, Tossa de Mar [Tossa; 41.7167°N, 2.9333°E], 29-VI to 19-VII-1968, Th. Palm, D. elongata det. L. Borowiec, MZLU [2005-03]; 1♂ diss., Totana [37.7667°N, -1.5°W], Murcia, V- 1938, Balaguer, D. elongata det. Petitpierre, GSWRL [2003-05]; 1♂ diss., 1♂, Valencia [39.4667°N, - 0.3667°W], Maru'der, MNMS [2003-02]; 1♂ diss., 2♀♀, Vera [37.2500°N, -1.8667°W], with Almira [Almeria], 4-VI-1977, brookside, D. elongata det. Steinhausen [1♂ diss.], D. elongata det. Eber 1985 [2♀♀], ZMHB [2006-02]; TUNISIA: 1♂ diss., Belvedere [36.82472°N, 10.18638°E], 25-IX-1929, A. Schatzmayer, [D. e. ab.] sublineata det. Burlini 1967, MSNM [2003-07]; 1♀ diss., [33.5378°N, 10.6761°E], 9- IV-1977, S. Mahunka, D. elongata det. V. Tomov, No. 99, HNHM [2003-08]; 1♂ diss., 1♀ diss., 1♂, 1♀, , 16.5 km north [33.6075°N, 9.005°E], 19-V-2000, A. Kirk, on Tamarix spp., GSWRL [2003-42, 2005- 50]; 1♂ diss., 1♀ diss., el Guetar [34.39°N, 8.84°E], [9.3 km southeast] Gafza [], 200m [elev.], Lortess, 18-IV-1993, R. Regalin, D. elongata det. D. Regalin 1995 [♀], Coll[ection] D. Sassi [♀], GSWRL [2003-61], MRSN [2003-08]; 1♂ diss., 1♀ diss., 2♀♀, Chott Sedjoumi [36.765°N, 10.15055°E], 27-IX-1929, A. Schatzmayer, D. e. ab. sublineata det. Burlini 1967, MSNM [2003-01, 03]; 1♂ diss., [Jedeida; 36.83111°N, 9.92416°E], 14-X-1929, A. Schatzmayer, [D. e. ab.] sublineata det. Burlini 1967, MSNM [2003- 05]; 2♂♂ diss., 1♂, 7♀♀, Marith, 18.7 km southeast on road C116, 33.5760°N, 10.4553°E, 20-V-2008, Javid Kashefi, on Tamarix sp. in desert wadi, shipment EBCLGR-JK-2008-004, New Mexico, Las Cruces, NMSU lab colony F4 generation, 8–27-X-2008, D. Guenther, voucher shipment GSWRL-2008-03, GSWRL [2008- 23, 24]; 1♀ diss., 2♀♀, Gabes [33.7425°N, 10.20567°E], Medenine road, 15-V-2000, A. Kirk, on Tamarix spp., GSWRL [2003-49]; 2♂♂ diss., Gabes, beach at Hotel Chems [33.20530°N, 11.20970°E], 6-V-2007, J. Kashefi, Tamarix sp. (not T. aphylla) (1♂), T. aphylla (1♂), shipment GSWRL-2007-01, GSWRL [2007-48, 49];1♂ diss., 3♂♂, 4♀♀, [33.70483°N, 8.942333°E], side, 18-V-2000, A. Kirk, on Tamarix spp., GSWRL [2003-43]; 2♂♂ diss., 4♀♀ diss., 1♂, 1♀, Sfax [34.7406°N, 10.7603°E], Gargour, Nahkia, 14- V-2000, A. Kirk, on Tamarix sp., GSWRL [2003-09, 12, 50, 65, 76, 79]; 4♂♂ diss., 4♀♀ diss., Sfax, 15 km south [34.66°N, 10.67°E; not as in some shipping records], 30-IX-2002, A. Kirk, on Tamarix spp., voucher USDA lab colony at Albany, California (22-I-2003, 2003, D. Bean), voucher USDA lab colony at Temple, Texas (1♂, 1♀, 26-VIII-2003, 5-IX-2003, L. Milbrath [nos. 1286, 1283]), shipment EIWRU-2002- 1008, GSWRL [2003-14, 15, 22, 23, 74, 75, 86, 87] [released in south Texas in 2005] [used in biological studies of Milbrath and DeLoach (2006a, 2006b), Milbrath et al. (2007), Herr et al. (in prep.), Bean and Keller (in prep.), and Thompson et al. (in prep.)]; 1♂ diss., Sfax, 15 km South [34.63980°N, 10.65280°E], 6-V-2007, J. Kashefi, Tamarix sp. (not T. aphylla), shipment GSWRL-2007-01, GSWRL [2007-50]; 1♀ diss., Sfax, 17 km South [34.65°N, 10.66°E], 6-III-1995, Kirk & Sobhian, on Tamarix spp., 9137, D. elongata det. I. Lopatin 1999, GSWRL [2004-15]; 1♂ diss., 2♂♂, Tozeur [33.92055°N, 8.13333°E], 14-XII-1928, Torre e Tasso, [D. e. ab.] sublineata det. Burlini 1967, MSNM [2003-06]; UNKNOWN COUNTRY: 1♂ diss., Pyrenneaen [not locatable, probably Pyrennes Mts., possibly France], Porrir, Scheider der Kelch, DEI [2003-12]; YEMEN: 2♀♀ diss., Yemen [specific locality not given], Arabia, Millingen [approx. location near Haraz Mts. (Millingen 1874): 14.87820°N, 43.65640°E], Fry Coll. 1908.100, BMNH [2003-04, 05]. Distribution. General. Diorhabda sublineata ranges from Portugal, Spain and France to Morocco, Senegal, Algeria, Tunisia, Egypt, Yemen, and Iraq (Map 4). Chatenet (2002) reported Diorhabda was little common in France and Petitpierre (1988) reported it as infrequent along the northeastern Mediterranean coast of Spain. It is apparently of sporadic occurrence in southern Spain, where surveys of Tamarix in July 2005 revealed no Diorhabda (J. Sanabria and J. Vela, pers. comm.). But Hopkins and Carruth (1954) report Diorhabda as common in Huelva Province of southwest Spain. Boehm (1908) found D. sublineata to be the most common galerucine in Egypt and Pierre Jolivet (Paris, France, pers. comm.) collected specimens which we have dissected from a dense population in the thousands at Ndiol Nar, Senegal. Reports of D. e. var.

78 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS sublineata (Weise 1890), D. e. ab. sublineata (Medvedev and Voronova 1977b, Medvedev 1982), and D. e. sublineata (Gressitt and Kimoto 1963a; Lopatin 1968, 1975, 1977b) from Mongolia or China should refer to D. carinulata. Further collections should reveal that the range of D. sublineata includes Western , , Libya, Sudan, Saudi Arabia, and Kuwait and yield specific locations in Portugal, north Yemen, and the Pontevedra (Galicia) and Huelva provinces of Spain. Tamarix aphylla and the probable host, T. nilotica, are found in east Africa in Ethiopia and Somalia (Baum 1978), where D. sublineata may also occur. Two Tamarix spp. are endemic to southern Africa (in , and ; Baum 1978), where surveys for D. sublineata should be made. Confirmed Records. We have dissected specimens from (see Material Examined) and confirmed the presence of D. sublineata (as D. e. var. sublineata, D. e. ab. sublineata, and D. sublineata) in the following countries with previous literature records (Map 4): Algeria (Lucas 1849, Peyerimhoff 1926), Egypt (Boehm 1908, Alfieri 1976), Tunisia (Normand 1937), and France (Laboissière 1934). New Records. We dissected specimens of D. sublineata from the following countries for which we find no previous specific reports of this species in the literature: Portugal, Spain, Morocco, Senegal, Iraq, and Yemen. With the exception of Yemen, we find past reports of D. elongata from these countries that should refer, at least in part, to D. sublineata (see above synonymy). Specimens at BMNH from Yemen with a label of “Millingen” were probably collected circa 1873 on a journey by Dr. Charles Millingen to north Yemen where he noted tamarisk in lowlands surrounding the Haraz Mountains (Millingen 1874). Unconfirmed Records. No country records of D. sublineata remain unconfirmed. On the basis of evidence discussed below, we accept as D. sublineata several locality records of Diorhabda from France, Spain, and North Africa. Diorhabda sublineata has been released in the United States (Texas), but establishment is not yet confirmed (Map 7, see Potential in Tamarisk Biological Control below for additional details). Specimens of D. sublineata were dissected from all 11 available locations from France. We examined a specimen of D. sublineata from France (IRSNB) with an identification label of D. e. var. carinata made by Laboissière in 1939. Therefore, we consider Laboissière’s (1934) report of D. e. var. carinata in France as D. sublineata, although D. elongata may also be present. Of 14 locations in Spain, D. sublineata was dissected from 13 of these while D. elongata was dissected from only one. Therefore, we believe Hopkins and Carruth (1954) found D. sublineata, rather than D. elongata, to be common in the Huelva Province of southwest Spain. Torres Sala (1962) reported D. sublineata (as D. elongata) from Comunidad Valenciana, Spain. Petitpierre (1988) found D. sublineata (as D. elongata) was infrequent along the northeastern coast of Spain. Specimens of D. sublineata were dissected from all 40 specific localities available from North Africa. D. elongata was also present at only one specific locality, Al Qahirah (Cairo), Egypt, and one general locality, Algeria, from a series predominated by D. sublineata. Consequently, we accept as D. sublineata all of the 23 unconfirmed North African collection records of D. sublineata (Lucas 1849; see also below Discussion- Taxonomy), D. e. ab. sublineata (Normand 1937, Alfieri 1976), D. e. ab. bipustulata (Normand 1937), and D. elongata (Peyerimhoff 1926; Normand 1937; Jolivet 1967; A. Kirk; USDA/ARS, Montferrier-sur-Lez, France, retired, pers. comm.; S. Doguet, pers. comm.). Weise (1925) reported that an expedition to Anglo- Egyptian Sudan collected D. sublineata (as D. elongata) from Halwan (= Heluan), Egypt, a location from which we dissected D. sublineata. This record was apparently incorrectly cited by Laboissière (1934) as a collection from Sudan, and this error was followed by Ogloblin (1936). Below are listed 25 unconfirmed locality records that we assign to D. sublineata (Map 4): ALGERIA: Hippône [Annaba; 36.9000°N, 7.7667°E] (Lucas 1849); Djamaa [33.5333°N, 6.0000°E], on Tamarix boveana (as T. bounopaea), III; Maison Carree [El Harrach; 36.7203°N, 3.14500E], on Tamarix africana, IX (Peyerimhoff 1926); Aurès Mountains, Beni Imloul Forest, Sidi Ali [=Ain Sidi Ali; 35.2833°N, 5.4667°E], 14-VI-1981, S. Doguet, on Tamarix; Biskra, Oued Beraze, 24-VI-1981, S. Doguet, on Tamarix; Guelma, Djebel Taya [36.507°N, 7.0828°E], 1-II-1968, S. Doguet, on Tamarix; Kabylie, El Adjiba [36.3258°N, 4.1503E], VI-1959, J.C. Bourdonné; Nemanchas Mountains, Khangat Sidi Nadji [Sidi Naji; 34.8167°N, 6.7000°E], 24-VI-1981, S. Doguet, on Tamarix (S. Doguet, pers. comm.); EGYPT: Asyut

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 79 [27.1828°N, 31.1828°E], IX; Cairo vicinity, V-X [D. sublineata dissected from same location]; Helwan [Halwan], wadis northeast, V (Alfieri 1976); Heluan [Halwan] [D. sublineata dissected from same location] (Weise 1925); Minya, IX [Al Minya; 28.1194°N, 30.7444°E]; Siala [Silah; 29.3561°N, 30.9689°E], VI [D. sublineata dissected from same location]; Sinnuris [29.4167°N, 30.8667°E], VI (Alfieri 1976, on Tamarix spp.); FRANCE: Narbonne [43.1833°N, 3.0000°], D. e. var. carinata (Laboissière 1934); MOROCCO: Essaouira, Ounara [31.5336°N, 9.5536°W], 3-V-1983, S. Doguet, on Tamarix; Tamelelt [Tamelelt Jdida; 31.8189°N, 7.5089°W], Marrekech [D. sublineata dissected from same location], 11-V-1983, S. Doguet, on Tamarix (S. Doguet, pers. comm.); Melilla area [33.3881°N, -7.14500°E], on Tamarix sp.; Sidi Mouna el Harati [Sidi Moussa el Harrati shrine; 34.0800°, -5.9600°E], on T. africana (Jolivet 1967); South Essaouira, Tamri [30.6780°N, 9.8253°W], 4-V-1983, S. Doguet, on Tamarix (S. Doguet, pers. comm.); SPAIN: Empúries [Ampurias; 42.1333°N, 3.1167°E] (Petitpierre 1988); Huelva [Province in southwest Spain; specific locality not given; app. locality: 37.25830°N, -6.9508°E], Tamarix gallica (Hopkins and Carruth 1954); Torroella de Montgri [42.0333°N, 3.1333°E]; Valls [41.2833°N, 1.2500°N] (Petitpierre 1988); TUNISIA: Le Kef [; 36.1822°N, 8.7147°E], Kairouan [35.6744°N, 10.1017°E], Medjez–el–Baba [Majaz al Bab; 36.6500°N, 9.6167°E] (Normand 1937); 1 spmn., Douz, 20.2 km N, 33° 37' 52" N, 8° 59' 58" E [33.6311°N, 8.9994°E], 19-V-2000, A. Kirk, Tamarix aphylla (Alan Kirk, USDA/ARS, Montpellier, France, retired, pers. comm.). D. sublineata × D. elongata Hybrid Morphology. All six studied adult male D. sublineata × D. elongata laboratory hybrid F1 and F2 progeny (Figs. 46–47) obtained from D.C. Thompson and B. Peterson (NMSU) can be distinguished from parental pure lines (Figs. 44–45) by a variety of anomalous hybrid genitalic character combinations in the endophallic sclerites. These diagnostic character combinations are not seen in laboratory pure lines of D. sublineata and D. elongata (Figs. 44–45) or in field collected material anywhere in the Palearctic, including areas of marginal sympatry for D. sublineata and D. elongata such as Portugal, Spain and Egypt (Figs.14, 16, 19, 21, 24, 26, 29, 31). Examples of anomalous character combinations in hybrids include: (1) length of blade (LB) of elongate endophallic sclerite (EES) is intermediate between the maximum length of blade for D. elongata (0.44 mm), and the minimum length of blade for D. sublineata (0.52 mm) (Figs. 46–48, NMSU 2006-1, 2); (2) length of spined portion (SL) of EES is intermediate between the maximum spined length for D. elongata (0.18 mm) and the minimum spined length for D. sublineata (0.38 mm) (Figs. 46–48, NMSU 2006-3, 4, 5); (3) length of blade of EES falls within the range for D. sublineata, but length of spined portion of the EES falls within the range for D. elongata (Figs. 47–48, NMSU 2006-6), or is intermediate between the species (NMSU 2006-3); (4) length of blade of EES falls within the range for D. sublineata, but connecting endophallic sclerite (CES) is absent as in D. elongata (Figs. 47–48, NMSU 2006-3, 4, 6); (5) palmate endophallic sclerite (PES) is broadly rounded with subdistal spines as in D. elongata, but length of blade of EES falls within the range of D. sublineata (Fig. 47—NMSU 2006-4); and (6) PES is truncate serrate with more than two spines being distal as in D. sublineata, but length of blade of EES falls within the range of D. elongata (Fig. 47—NMSU 2006-5). All hybrids examined lack the CES (Figs. 46–47) which is indicative of D. sublineata (Fig. 45). Of the six D. sublineata/D. elongata hybrids illustrated, most bear the external coloration and pattern of elytral vittae found in D. elongata, and only one hybrid (Fig. 47—NMSU 2006-5) had submarginal and subsutural elytral vittae extending into the base of the elytra as is sometimes found in D. sublineata. The morphologies of later generation hybrids or various types of backcross hybrids are not characterized. Many of the male D. sublineata × D. elongata hybrids possess genitalic abnormalities in the form of conspicuous abnormal sclerites (AS) of varying number, size, shape and location on the endophallus (Figs. 46–47). Abnormal sclerites are not seen or are small and inconspicuous in parental pure lines (Fig. 44–45). Externally normal appearance with genitalic abnormalities is commonly observed in interspecific hybrids between some sibling species of Drosophila fruit flies (e.g., Hollocher et al. 2000). Abnormal endophallic sclerites in D. sublineata × D. elongata hybrids are evidence of genetic incompatibilities and possibly some level of reduced hybrid fitness and postzygotic reproductive isolation.

80 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Discussion. Taxonomy. Galeruca sublineata Lucas (1849) (as Galleruca sublineata) was described from Hippône (= Annaba), Algeria, and synonymyzed under G. elongata by Reiche and Saulcy (1858). Weise (1890) proposed D. elongata var. sublineata as extending in range from North Africa to Mongolia. Weise (1924) later proposed the aberration D. e. ab. sublineata and this has been followed by several taxonomists (Ogloblin 1936, Normand 1937, Alfieri 1976, Warchalowski 2003). Chen (1961) excluded D. e. ab. sublineata from Mongolia and included the Mongolian population under the new subspecies D. e. deserticola described from western China. Apparently unaware of Chen’s designation, Gressit and Kimoto (1963) proposed the subspecies D. e. sublineata as occurring from Mongolia and western China west to North Africa. The catalogue of Lopatin et al. (2004) lists D. e. sublineata as a junior synonym of D. e. elongata. In order to characterize the genitalia of G. sublineata, we examined specimens that matched the species description of G. sublineata from Algeria and Tunisia, in the broad vicinity of the type locality of Annaba, Algeria. In the color habitus illustration accompanying the species description for G. sublineata by Lucas (1849, Plate 44, Fig.8), the submarginal and subsutural elytral vittae extend well into the basal half of the elytra (as in Fig. 5). This characteristic of the elytral vittae can be used to distinguish G. s ub l in ea t a from D. elongata. In D. elongata, the elytral vittae are often absent, as in the color habitus illustration with the species description by Brullé (1832, Plate 44, Fig. 10), and, when present, the elytral vittae are confined to the apical half of the elytra (Fig. 1, see D. elongata - Discussion-Taxonomy above). From 12 locations in Algeria and Tunisia, we selected ten male and seven female specimens matching the description of G. sublineata with elytral vittae extending well into the basal half of the elytra. We are confident that these 17 specimens represent G. sublineata as described by Lucas (1849). We studied the genitalia in the 17 above specimens matching G. sublineata from Algeria and Tunisia. In the ten males, the endophalli bear a combination of five characteristics distinguishing them from those of D. elongata (characterized above) and the holotypes of D. carinulata, D. meridionalis,and D. carinata (Figs. 18–19 of Berti and Rapilly 1973) (see Male-Genitalia above; Figs. 14–33). In the seven females matching G. sublineata, the vaginal palpi were triangulate and narrowly rounded, as in D. carinata, and distinct from the broadly rounded vaginal palpi of D. elongata. The distinct male and female genitalic characters of specimens matching G. sublineata are strong evidence of reproductive isolation from other species in the group, and especially from the two species with which it is marginally sympatric, D. elongata and D. carinata (see Biogeography below; Map 1, Table 8). Therefore, we remove G. s u bl in e at a from synonymy with D. elongata and restore the species D. sublineata (Lucas) REVISED STATUS. Although D. sublineata is allopatric with D. carinulata and D. meridionalis, their status as reproductively isolated is supported by the number of diagnostic genitalic characters separating D. sublineata from D. carinulata and D. meridionalis (6–8 characters) being greater than the number of characters separating the moderately sympatric D. carinata from D. carinulata (5 characters) (Table 5). Further evidence for reproductive isolation between D. sublineata and D. elongata is also found in previously discussed differences in component ratios of putative aggregation pheromones and reduced F2 hybrid egg viability. The elytral vittae of D. sublineata can resemble that of D. elongata in being confined to the apical half of the elytra or entirely lacking. The color habitus illustration of “D. elongata” from southern Europe in Plate 29, Figure 7 of Chatenet (2002), with the elytral striping extending through the entire basal half of the elytra, is actually of a specimen of D. sublineata. If D. sublineata interbreeds as a subspecies with D. elongata where these species are marginally sympatric in Portugal, Spain and Egypt, we would expect intermediate hybrid forms to be evident in areas of range contact as seen in other chrysomelid subspecies, such as Diabrotica virgifera virgifera and D. v. zeae Krysan and Smith (Krysan et al. 1980). For example, we would expect hybrid characteristics such as varying degrees of development of the connecting endophallic sclerite and the lateral appendage of the palmate endophallic sclerite and intermediate lengths in the blade of the elongate endophallic sclerite. Diagnostic hybrid characteristics are found in examined laboratory produced D. sublineata × D. elongata hybrids (Figs. 46–47; see D. sublineata × D. elongata Hybrid Morphology above). However, we find no specimens with diagnostic hybrid characteristics in field collections of 157 male D. elongata and 89 male D. sublineata. These include 37 specimens from the areas of range overlap; 6 male D. elongata in Egypt, (Figs. 19, 24,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 81 29—Cairo), Portugal (Fig. 19—Portugal), and Algeria (Fig. 19—Algeria) and 42 male D. sublineata in Egypt (Figs. 21, 26, 31—Cairo), Algeria (Fig. 21—Algiers), Spain (Fig. 21—Valencia), and Portugal. The lack of intermediate hybrid forms in areas of sympatry is firm evidence of reproductive isolation between D. elongata and D. sublineata, and we are confident that D. sublineata is not a subspecies of D. elongata. The diagnostic characters separating D. sublineata from D. elongata comprise three discrete and two near discrete male genitalic characters and one discrete female genitalic character for a total of six diagnostic genitalic characters, four of which are discrete (see keys and Table 5). In comparison, a total of five discrete genitalic characters are diagnostic in distinguishing the moderately sympatric species D. carinata and D. carinulata (Table 5). Even if D. elongata and D. sublineata were allopatric rather than marginally sympatric, their degree of divergence in genitalic characters is similar to that found in the related sympatric species pair D. carinata/D. carinulata (see also D. elongata group Stenophenetic Analysis, Fig. 49), further justifying their status as separate species (see Helbig et al. 2002). As noted with other members of the D. elongata group, we find that the external characters that have been used in separating D. sublineata from other sibling species are inadequate and that genitalic characters must be used. Weise (1890) proposed that two sharply defined stripes on the elytra (elytral vittae) that joined apically on D. sublineata (as D. e. var. sublineata) is a distinguishing character from D. elongata. The use of this character as diagnostic has been followed by Laboissière (1934) and Warchalowski (2003). However, apically joined elytral vittae variably occur in both D. sublineata and D. elongata. Chen (1961) noted that the ventral tarsal pubescence was generally distributed in D. e. ab. sublineata, as opposed to being medially absent (leaving a median glabrous area) in D. e. deserticola. However, we found that the pattern of ventral tarsal pubescence in specimens of D. sublineata from North Africa is the same as that described by Chen for D. e. deserticola, and regard this character as unsuitable for taxonomic diagnosis. We have dissected specimens of D. sublineata that were misidentified by taxonomists using external diagnostic characters as D. elongata from five countries and D. e. var. carinata from Algeria (see Material Examined). We examined four specimens (from ZMHB) with old hand-written determination labels of D. e. var. sublineata from Weise’s collection that were collected by Potanin from Mongolia. All four of these specimens are D. carinulata. These specimens are probably from the same series listed by Weise (1890) as D. e. var. sublineata and collected by G.N. Potanin from central Mongolia. All material we have examined from Mongolia and China are conspecific with D. carinulata (discussed below), and D. sublineata is absent from these areas, the easternmost occurrence of D. sublineata being in Baghdad, Iraq (Map 1). Normand (1937) described the aberration D. elongata ab. bipustulata Normand from Kairouan, Tunisia in the area where D. sublineata is common (Maps 1 and 4). This aberration is distinguished by two dark spots near the base of the elytra on either side of the scutellum. We saw this uncommon color variant in one specimen (from Morocco) of over 150 examined specimens of D. sublineata. It was also seen in six specimens (from Greece and Cyprus) of 200 examined specimens of D. elongata, two specimens (from Ash Sharqat, Iraq) of over 150 examined D. carinata, and one specimen (from 74–164 km NW Turpan, China) of over 300 examined D. carinulata. Diorhabda elongata apparently is rare in Algeria and probably rarely occurs, if at all, in Tunisia, from which all 17 dissected specimens from all 12 available locations were D. sublineata. Therefore, we consider D. e. ab. bipustulata as a junior synonym of D. sublineata NEW SYNONYMY. Diorhabda sublineata and D. carinata are the most morphologically similar species in the D. elongata group, and we considered the possibility that D. sublineata might be a subspecies of the earlier described D. carinata. The only character for consistent separation of D. sublineata from D. carinata is the presence of the connecting endophallic sclerite in the male (Figs. 16, 21, 26, 31). This connecting sclerite is not a trivial character in that it influences the three dimensional configuration of the inflated endophallus. In D. sublineata killed and preserved while copulating, the inflated endophallus is bent at the connecting sclerite and this is not seen in inflated endophalli of D. carinata. If D. sublineata and D. carinata were interbreeding subspecies, we should have observed intermediate forms in the critical distinguishing morphological character of the presence or absence of the connecting endophallic sclerite. Intermediate forms should increase along a

82 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS geographic cline approaching the point of range contact in Iraq. Specifically, we should have observed a transition from the fully developed connecting endophallic sclerite of D. sublineata to increasing incidence of faint lines in place of a connecting endophallic sclerite, to the total lack of connecting endophallic sclerite found in D. carinata. From Baghdad, Iraq, we examined a single male D. sublineata with a normal connecting endophallic sclerite (Fig. 21 – Baghdad) and 6 male D. carinata, none of which bore even a faint line where the connecting endophallic sclerite would be found (Fig. 20—Baghdad). The lack of intermediate forms in field collected material of 98 male of D. carinata and 69 male of D. sublineata, is evidence of reproductive isolation. Neither males nor females of D. sublineata and D. carinata were more difficult to identify to species near the contact zone of Iraq as would be expected were these to be interbreeding subspecies. Previously discussed differing component ratios in the putative aggregation pheromones of D. carinata and D. sublineata and the contrasting results of crossing each of these species with D. elongata are additional evidence of reproductive isolation. Common Name. The vernacular name “subtropical tamarisk beetle” refers to most of the main distribution of D. sublineata (31–38°N; Fig. 51B) falling within the subtropical climates of North Africa (region from ca. 23.5–35°N with mild winters). This subtropical distribution contrasts with that of the marginally sympatric sibling species, D. elongata, which has its main distribution (37–41°N) entirely north of 35°N and which is rarely found below 35°N (Fig. 51B). Diorhabda sublineata has the strongest presence of any Diorhabda sibling species in subtropical to tropical biomes, including the Flooded Grasslands and Savannas and the Tropical and Subtropical Grasslands, Savannas and Shrublands (Table 9; Fig. 52B). Biology. Host Plants. Diorhabda sublineata (as D. elongata) is reported from T. africana Poiret and T. boveana Bunge (as T. bounopaea Gay) in Algeria (Peyerimhoff 1926) and T. africana and Tamarix sp. in Morocco (Jolivet 1967) (Table 1). Reports of T. gallica as a host of D. elongata in France (Laboissière 1934) and Spain (Huelva Province, Hopkins and Carruth 1954) should also refer to D. sublineata. de Candolle is a new host record from collections in Ndiol Nar, Senegal by P. Jolivet (BMNH collection). Tamarix aphylla is a new host for D. sublineata (as D. elongata) derived from a single adult collected by A. Kirk (pers. comm.) 20.2 km north of Douz, Tunisia. We examined other specimens of D. sublineata that Kirk collected from Tamarix sp. (not T. aphylla) at six locations in Tunisia. Diorhabda sublineata is reported from Tamarix spp. in Egypt (Boehm 1908, Alfieri 1976) and has been found in several locations along the River where T. mannifera (Ehrenberg) Bunge and T. nilotica (Ehrenberg) Bunge, close relatives of T. senegalensis (Baum 1978) are common, and these species may also serve as hosts. The center of distribution of T. canariensis is in Algeria, Morocco and Spain (Baum 1978), and it is another likely host (Map 4). Tamarix arabica Bunge occurs in Yemen (Baum 1978), where it could serve as a host of D. sublineata. No-choice larval host suitability studies by Milbrath and DeLoach (2006a) confirm that D. sublineata larvae from Tunisia can survive to adulthood only on plants of the order Tamaricales, including Tamarix (Tamaricaceae) and, to a significantly lesser degree, on three North American Frankenia spp. (Frankeniaceae): F. s a li na , F. johnstonii, and F. ja m e si i. In field cage no-choice studies, oviposition by D. sublineata on F. ja m e s ii and F. johnstonii was not different from that on non-host coyote willow (Salix exigua Nutall) and adults experienced increased mortality compared to T. ramosissima × T. chinensis treatments (Milbrath and DeLoach 2006a). Multiple-choice adult oviposition studies in field cages (Milbrath and DeLoach 2006a) revealed that the three North American Frankenia spp. provide a little attraction for oviposition compared to Tamarix. However, surveys by Alan Kirk in Tunisia of May 2000 revealed D. sublineata (as D. elongata) on Tamarix spp. from five of eight collection sites, but no Diorhabda were found on Frankenia at six collection sites in areas near Tamarix, even where Frankenia was found adjacent to Tamarix on which D. sublineata was abundant (DeLoach et al. 2003b; A. Kirk, pers. comm.). In field cage multiple-choice studies, D. sublineata oviposited significantly less on T. aphylla than on most of the invasive North American tamarisks, including T. ramosissima, T. ramosissima × T. chinensis, T. ramosissima × T. canariensis/T. gallica, T. chinensis × T. canariensis/T. gallica, and T. parviflora (Milbrath and DeLoach 2006a, Milbrath and DeLoach 2006b). Oviposition did not differ between T. aphylla and T. canariensis/T.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 83 gallica in one of these field cage tests (Milbrath and DeLoach 2006b) while oviposition was significantly lower on T. aphylla than on T. canariensis/T. gallica in the other test (Milbrath and DeLoach 2006a). For oviposition, T. aphylla is accepted by D. sublineata to the same degree as T. ramosissima × T. chinensis in no- choice field cage studies (Milbrath and DeLoach 2006b). Tamarix aphylla is at moderate risk to damage by D. sublineata in the field and the degree to which D. sublineata would damage T. aphylla is difficult to predict, especially in the absence of other Tamarix spp. (Milbrath and DeLoach 2006b). Moran et al. (in press) found that D. sublineata (ex: Sfax, Tunisia) × D. elongata (ex: Sfakaki, Crete, Greece) hybrids demonstrate a clear preference to T. canariensis/T. gallica over T. aphylla in open-field tests near Kingsville, Texas. Frankenia is at very low risk of damage from D. sublineata (Milbrath and DeLoach 2006a). Risk of damage to both T. aphylla and Frankenia by D. sublineata is probably much lower when these plants are not in the proximity of preferred Tamarix spp. (e.g., Blossey et al. 2001). Ecology and Phenology. We found no reports on the ecology and phenology of D. sublineata in the Palearctic. Adult collection dates in the literature and our examined material are from January to November in Egypt, 22 January to 14 December in Tunisia (both from examined material), January to September in Algeria (Peyerimhoff 1926, Doguet, pers. comm.) and April to October in France and Spain (this study). From these data, adults are found almost year round in North Africa. Milbrath et al. (2007) found that D. sublineata (as D. elongata from Sfax, Tunisia) overwintering at Temple, Texas had ca. 92% survival from early November through the beginning of March but survival dropped to ca. 62–67% by mid-March when the tamarisk leaves were just budding. Overwintered adults began ovipositing in late March giving rise to five generations and a partial sixth generation. Fifth generation adults emerging in early September oviposited for several weeks before ceasing oviposition in November when they appeared to enter diapause. Development and Reproduction. Milbrath et al. (2007) found that, at 28°C, D. sublineata (as D. elongata from Sfax, Tunisia) had a development time of 18.6 days from egg to adult (with 89% survival), a fecundity of 208 eggs, and a population doubling time of 5.5 days. These values were all very similar to those found for D. carinulata (Turpan and Fukang), D. elongata (Crete), and D. carinata (Uzbekistan) (all as D. elongata) in the same study. Natural Enemies. Nosema sp. microsporidians infected D. sublineata larvae and adults collected southeast of Marith, Tunisia in 2008 (shipment EBCLGR-JK-2008-004) (D. Bean, pers. comm.). Biogeography. Comparative. Diorhabda sublineata differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) strongly maritime and generally found within ca. 200 km from a sea coast, usually below 300 m elevation (but can range to 2,600 m elevation); (2) favors subtropical Mediterranean woodlands or Flooded Grasslands, Savannas and Shrublands biomes; and (3) latitudinal range of 16–44°N and most commonly collected from 31–38°N (Table 7; Figs. 51–52). In contrast, the marginally sympatric D. elongata has a southern range limited to 30°N and it is much less common south of 35°N. In addition, D. elongata is rare or lacking in flooded grassland and desert biomes in which D. sublineata is found. Although both D. elongata and D. sublineata share T. gallica as one of their hosts (Table 1), we have seen no mixtures of D. sublineata and D. elongata within series collected from specific localities such as are found with D. carinata and D. carinulata, or D. carinata and D. meridionalis. This may be the result of D. sublineata dominating in areas where it is marginally sympatric with D. elongata in Spain, Algeria, Egypt and probably elsewhere along the Mediterranean coast of North Africa (Map 1, Table 8; see also discussion under Biogeography for D. elongata). The possibility that D. sublineata may occur in Italy where it may be rare and dominated by D. elongata should be investigated. Diorhabda sublineata is marginally sympatric with D. carinata in Baghdad, Iraq, where their ranges meet, and it is allopatric with D. carinulata (Map1; Table 8). Both D. carinata and D. carinulata differ from D. sublineata in being primarily continental in distribution, mostly found in desert and grassland biomes rather than the Mediterranean biome, and being common north of 38°N (Figs. 51–52). Diorhabda sublineata is also allopatric with D. meridionalis (Map 1; Table 8). Diorhabda meridionalis differs from D. sublineata in

84 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS being more commonly collected further south at 26–30°N, and in preferring the desert biome over the Mediterranean biome (Figs. 51–52). Descriptive. Diorhabda sublineata primarily inhabits the southwestern Palearctic realm and is the only member of the D. elongata group known to occur in the Afrotropical (in Senegal and Yemen) (Map 1). It has most commonly been collected from the Mediterranean Forests, Woodlands and Scrub biome from 31–38°N (Fig. 52B). The next most important biome for D. sublineata is the Flooded Grasslands and Savannas biome from ca. 24–34°N (Map 4; Fig. 42B). Diorhabda sublineata is also found in two additional biomes: the Deserts and Xeric Shrublands biome from 25–33°N in North Africa and Iraq (at ca. 20–600 m elevation), and the Montane Grasslands and Shrublands biome to ca. 2,600 m elevation at Tacheddirt, Morocco (latitude 31°N) (Map 4; Table 7). Diorhabda sublineata occurs primarily in three ecoregions of the Mediterranean Forests, Woodlands and Scrub biome (Olson and Dinerstein 2002), based on frequency of collections (Map 4): Mediterranean Woodlands and Forests of coastal Morocco, Algeria and Tunisia (at ca. 0 to 1,700 m), Mediterranean Acacia- Argania Dry Woodlands and Succulent Thickets of Morocco (at ca. 0 to 600 m), and Mediterranean Dry Woodlands and Steppe of Algeria and Tunisia (at ca.0 to 600 m). Primary ecoregions of D. sublineata in the Flooded Grassland and biome are the Nile Delta Flooded Savanna of Egypt (at ca. 0 to 80 m elevation) and Saharan Halophytics of Algeria and Tunisia (at ca. 20 to 120 m elevation). Diorhabda sublineata was the most common galerucine reported in Egypt in 1908 (Boehm 1908). Diorhabda sublineata is found only in the Mediterranean Forests, Woodlands and Scrub biome in the northern portion of its range at ca. 37–42°N in France and Spain (Map 4). Diorhabda sublineata (as D. elongata) is common but not damaging in parts of southern Spain (Hopkins and Carruth 1954) and it is uncommon along Mediterranean coast of northeastern Spain (Petitpierre 1988) and southern France (Laboissière 1934, Chatenet 2002). We found few reports of D. sublineata in the southern portion of its range from 16–25°N in Africa and the Arabian Peninsula (Map 4). Additional collection efforts may reveal it to be more common in these areas. P. Jolivet (pers. comm.) reported August populations in Senegal reaching the thousands where he collected material that we examined from the Tropical and Subtropical Grasslands, Savannas and Shrublands biome. The northwestern portion of the range of D. sublineata (France, Spain, Morocco, Algeria and Tunisia) closely follows that of known host T. boveana and the suspected host T. canariensis (Map 4). Both hosts T. gallica and T. africana widely overlap the northwestern range of D. sublineata, but extend further east into Italy from which D. sublineata is not known. In Egypt, the distribution of T. nilotica and T. mannifera along the Nile River (Baum 1978) corresponds well with that of D. sublineata. Potential in Tamarisk Biological Control. Summary. The subtropical tamarisk beetle can potentially be effective in biological control of T. ramosissima/T. chinensis and hybrids with T. canariensis/T. gallica in the extreme southwestern U.S. Diorhabda sublineata may be the most suitable Diorhabda species for introduction into the Mediterranean biome below 37°N in California and the southern Chihuahuan Desert in Mexico (Map 13) (including the Mapimian and Saladan regions of Map 6 in Morafka [1977]). In the absence of D. meridionalis, D. sublineata is probably the most suitable species for a major portion of maritime subtropical deserts such as the Sonoran Desert and Tamaulipan Mezquital xeric shrubland. Discussion. We find no anecdotal reports of the subtropical tamarisk beetle damaging Tamarix in the Old World, but this species can be locally abundant in places such as Senegal and Kom Ombo, Egypt, from which a large series of 106 specimens was collected in November. Tamarix ramosissima and its relatives are not recorded hosts of D. sublineata. Hybrids of T. ramosissima/T. chinensis with T. canariensis/T. gallica are common in Texas (J. Gaskin, pers. comm.), and D. sublineata may prefer these hybrids more than would other Diorhabda. Diorhabda sublineata has a moderate risk of damaging T. aphylla (Milbrath and DeLoach 2006b) and very low risk of damaging Frankenia (Milbrath and DeLoach 2006a), and both these risks are probably much reduced at less proximity to preferred Tamarix spp. (e.g., Blossey et al. 2001). The subtropical tamarisk beetle is most commonly found in the Mediterranean Forests, Woodlands and Scrub biome at 31–38°N (Map 4, see Biogeography). In North America, Mediterranean ecoregions

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 85 corresponding to this area include the California Coastal Sage and Chaparral ecoregion of California and Baja California, the California Montane Chaparral and Woodlands and southern portions of the California Interior Chaparral and Woodlands (Map 13). Additional North America habitats should be similar to where D. sublineata is found in the subtropical maritime Flooded Grassland and Savannah biome from 24–34°N in North Africa that are surrounded by the subtropical maritime Deserts and Xeric Shrublands biome. In North America, this habitat particularly includes lower elevation (below 300m) portions of maritime subtropical Deserts and Xeric Shrublands biome from ca. 24–35°N in the following ecoregions: southern Mojave Desert, Sonoran Desert, Tamaulipan Mezquital along the lower Rio Grande, and Tamaulipan Mattoral (Map 13). Along the Rio Grande near Presidio in the Big Bend region, the distribution of the Rio Grande cottonwood (P. deltoides subsp. wislizenii) more common in the Trans-Pecos Chihuahuan transitions to the more southern Chihuahuan desert species, Meseta cottonwood (Populus fremontii subsp. mesetae) (Powell 1998). Correspondingly, the Big Bend area near Presidio may correspond with a transition zone for more northern desert D. carinata and D. carinulata into more southern desert D. sublineata. Morafka (Map 6 of Morafka 1977) mapped a transition from northern Trans-Pecos Chihuahuan Desert to more southern Mapimian Chihuahuan Desert at approximately the interface of our predicted ranges of D. carinata/D. carinulata and D. sublineata in the Big Bend region (Map 13). Tamarix aphylla is not a primary host for D. sublineata, but it is an indicator of the subtropical climate in North Africa where D. sublineata is most common (Map 4), and the most suitable areas in North America for D. sublineata may also occur within the area warm enough for T. aphylla to flourish (see Map 7). In the absence of D. meridionalis, D. sublineata would be the most suitable species for all of the Tamaulipan Mezquital in Texas according to our HSI model. Diorhabda sublineata appears to be well adapted to more southern latitudes in terms of initiation of diapause at shorter photoperiods (later in season) in the south (Bean and Keller in prep.) and can have high overwintering survival (ca. 65%) at Temple (Milbrath et al. 2007). Species distribution models incorporating climatic data are in preparation to better predict habitat suitability. Efforts to establish populations of D. sublineata from near Sfax, Tunisia near Ricardo, Texas on T. canariensis/T. gallica were not successful, possibly partly due to the small isolated poorly vigorous stand of tamarisk at this site. Plans are being made to release D. sublineata from near Marith, Tunisia in 2009 at better quality release sites with larger and more vigorous T. chinensis/T. canariensis stands along the Rio Grande and tributary streams in west and south Texas, such as at Ruidosa. Releases of any Diorhabda in southern California are awaiting clearance with USDA-APHIS (T. Dudley, pers comm.).

Diorhabda carinulata (Desbrochers, 1870) northern tamarisk beetle (Figs. 7, 8, 17, 22, 27, 32, 37, 42)

Galeruca carinulata Desbrochers, 1870:134 (Type locality: Sarepta, Russia). Diorhabda elongata var. sublineata: Weise, 1890:484 (part, central Mongolia). Diorhabda elongata var. carinata: Weise, 1893:635 (part, Sarepta, Russia). Diorhabda elongata ab. carinata: Weise, 1924:78 (part; world catalog; Sarepta, Russia); Winkler, 1924–1932:1307 (part; Palearctic catalog; southern Russia) Diorhabda elongata ab. sublineata: Weise, 1924:78 (part; world catalog; Mongolia); Laboissière, 1934:54 (part, Mongo- lia); Winkler, 1924–1932:1307 (part, Palearctic catalog, Mongolia); Medvedev and Voronova, 1977b:238 (part, keys, Mongolia); Medvedev, 1982:259 (part, keys to adults and larvae, Mongolia). Diorhabda koltzei ab. basicornis Laboissière, 1935:324 (Type locality: Khotan, Taklamakan Desert, Turkestan [Hotan, Xinjiang Uygur Zizhiqu, China]) (NEW SYNONYM); Wilcox 1971:63 (world catalog, Karakorum [western China]). Diorhabda elongata: Ogloblin, 1936:79 (part, Russia, transcaucasus, Iran, central Asia, central Mongolia); Rusanov, 1949:118 (part, Central Asia, as Diorrhabda); Kyrzhanovskiy, 1952:198 (part, Turkmenistan), 1965:392 (part, mid- dle Asia); Pavlovskii and Shtakelberg, 1955:566 (part, southwest Russia, transcaucasus, central Asia, Iran, Mongo- lia, as Diorrhabda); Yakhontov and Davletshina, 1955:58 (part, biology, Amu Darya Delta, northern Uzbekistan); Sinadsky, 1957:950, 1963:84, 1968:64 (part, biology, Amu Darya Valley, Uzbekistan, as Diorrhabda); Yakhontov,

86 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS 1959:338 (part, biology, Uzbekistan); Medvedev, 1959:118 (part, Turkmenistan); Kulinich, 1962:73 (part, biology, Tajikistan); Kryzhanovsky, 1965:392 (part, middle Asia); Pripisnova, 1965:83 (part, biology, Tajikistan); Kulenova 1968:171 (part, southeastern Kazakhstan); Lopatin and Tadzhibaev, 1972:591 (part, Tajikistan); Lopatin, 1977a:282 (part, keys, Asia); Medvedev and Voronova, 1977a:339 (Mongolia), 1977b:238 (part, keys, Mongolia), 1979:119 (catalog for Mongolia, hosts); Davletshina et al., 1979:79 (part, southwest Kyzyl–Kum Desert, central Uzbekistan); Ogloblin and Medvedev, 1971:83, 89 (part, key to larva, European Russia); Medvedev, 1982:99 (part, keys to adults and larvae, Mongolia); Samedov and Mirzoeva, 1985:712 (part, Azerbaijan); Lopatin and Kulenova, 1986:129 (part, Kazakhstan); Kovalev, 1995:78 (part, south central Palearctic); Myartseva, 1995:4, 1999:1; 2001:1 (part, biology, Turkmenistan); DeLoach et al., 1996:253 (part, China); Ishkov, 1996:30 (part, Ili River Valley, Kazakhstan); White, 1996:392 (part, China); Mityaev and Jashenko, 1997:4, 1998:7, 1999:6, 2000:14, 2001:9, 2007:3 (part, biology, ecology; southeast Kazakhstan); Gruev and Tomov, 1998:70 (part, Transcaucasus, mid-Asia, Mongolia); USDA- APHIS, 1999:13395 (Fukang, China; approval of initial introduction into U.S.); Anonymous, 2001:52N (part, Kazakhstan, China); Gould and DeLoach, 2002:302 (part, biological control, North America); Dudley et al. 2000:346 (part, biological control, North America), 2001:260 (part, biological control, North America); Stenquist, 2000:492 (Kazakhstan); Jolivet, 2001:134 (part, U.S. introduction); Vail et al. 2001:37 (part, U.S. introduction); DeLoach et al., 2003a:230 (part, China, Kazakhstan), 2003b:126 (part, host range; Uzbekistan, Turkmenistan, Iran); Khamraev, 2003:11 (part, Uzbekistan); Petroski, 2003:3234 (synthesis of aggregation pheromone components); Mil- brath et al., 2003:225 (part, China, Kazakhstan); Riley et al., 2003:69,189 (part, catalog, North America [intro- duced]); Warchalowski, 2003:328 (part, taxonomic keys, Caucasus, Central Asia, and Mongolia); Bieńkowski, 2004:76 (part, keys, eastern Europe (part, southern Russia, Caucasus, Iran, Iraq, Central Asia, Kazakhstan, Mongo- lia); DeLoach and Carruthers, 2004b:311 (part, China and Kazakhstan); Dudley and DeLoach, 2004:1542 (part, bio- logical control, North America); Lopatin et al., 2004:127 (part, central Asia); Cossé et al., 2005:657 (aggregation pheromone, Fukang, China, population), 2007:2695 (Fukang, China; attraction to green leaf volatiles); Dudley, 2005a:13, 2005b:42N (part, biological control, ex: China and Kazakhstan); Herrera et al., 2005:775 (part, develop- mental rates, Fukang, China, population); Carruthers et al. 2006:71, 2008:258 (part, biological control, ex: China and Kazakhstan); Milbrath and DeLoach, 2006a:32 (part, host specificity, Turpan, China, populations); Geraci et al. (in press) (remote sensing of defoliation, Nevada); Milbrath et al. (2007) (part, biology, Fukang, China, population); Dudley et al., 2006:137 (part, releases in Nevada, Fukang, China; host range, Fukang and Turpan, China); Longland and Dudley (2008) (ecology, North America); Dudley et al. (in prep.) (host range, Nevada); Thompson et al. (in prep.) (part, laboratory hybridization, Fukang, China). Diorhabda rybakowi: Mityaev, 1958:86 (part, biology, Kazakhstan, as rybakovi). Diorhabda elongata deserticola Chen, 1961 (Type locality: Yuli (= Wei–li), Xinjiang Uygur Zizhiqu, China) (NEW SYNONYM); Gressitt and Kimoto, 1963b:930 (China); Wilcox, 1971:63 (world catalog, Xinjiang Uygur Zizhiqu, China); Tian et al., 1988:24 (biology, ecology; Ningxia Province, China); Bao, 1989:45 (biology and control as pest; Nei Mongol Zizhiqu, China); Xiao, 1992:537 (biology, China); Sha and Yibulayin, 1993:7 (biology, ecology, control as pest, China, Xinjiang Prov.); Chen et al., 2000:44 (control as pest; Alashan Meng, Nei Mongol Zizhiqu, China); USDA-APHIS, 2005:1 (Fukang, China; large-scale introduction to 13 states in northwestern U.S.); Li et al., 2000:48 (biology, ecology; China, Xinjiang Prov., Fukang); Eberts et al., 2001:3 (Fukang, China, establishment in Colorado); Lair and Eberts, 2001:3 (China, potential introduction into north Texas); Zhang 2002:1 (biology and ecology, Xinji- ang Uygur Zizhiqu, China); DeLoach et al., 2003a:229 (China, Kazakhstan), 2003b:117 (host specificity; Turpan and Fukang, China and Chilik, Kazakhstan; host range; China, Kazakhstan, Mongolia), 2004:505 (field establish- ment in North America from populations in Fukang, China and Shelek, Kazakhstan), (2008, in prep.) (part, China); Lewis et al., 2003a:148 (host specificity; Fukang, China and Chilik, Kazakhstan), 2003b:101 (developmental and reproductive biology; Fukang, China and Chilik, Kazakhstan); Meng and Baoping, 2003:99, 2005:192 (biological control; Xinjiang Uygur Zizhiqu, China), 2006:189 (ecology; Xinjiang Uygur Zizhiqu, China); DeLoach and Carru- thers, 2004a:10 (field establishment in North America from populations in Fukang, China and Shelek, Kazakhstan); Ding, 2004:57 (control as pest; Anxi County, Gansu Sheng, China); Ming et al. 2004:283 (Xinjiang Uygur Zizhiqu, China); Lopatin et al., 2004:127 (east Kazakhstan, northwest China); Aber et al., 2005:63 (field monitoring in Colo- rado; ex: Fukang, China); Dudley and Kazmer, 2005:265 (host specificity, Fukang, China, population); Peng et al., 2005:63 (control as pest; Gansu Sheng, China); Zheng et al., 2005:136 (China); Ding et al., 2006:1442 (biological control, ex: Kazakhstan, China); Li and Wang, 2006:27 (biology, control as pest; Gansu Sheng, China); Bean et al., 2007a:15 (diapause, Fukang, China); Mityaev and Jashenko, 2007:8 (biological control, ex: Kazakhstan, China); Bean et al., 2007b:531 (diapause, Fukang, China); Hudgeons et al., 2007a:158 (biological control; Fukang, China, Chilik, Kazakhstan), 2007b: 215 (part, tamarisk damage in Nevada, ex: Fukang, China); Dalin et al. (in press) (host range; Turpan, China). Diorhabda elongata sublineata: Gressitt and Kimoto, 1963a:407 (part; keys; northwestern China and Mongolia); Lopa- tin, 1968:214 (Mongolia); 1970:254 (Mongolia, on Tamarix); 1975:219 (Mongolia); 1977b:154 (Mongolia). Diorhabda carinulata carinulata: Berti and Rapilly, 1973:881 (restored species; Sarepta, Russia); DeLoach et al., 2003b:126; Warchalowski, 2003:328 (taxonomic keys, southern Russia).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 87 Diorhabda deserticola: Yu et al., 1996:94 (taxonomic keys and descriptions, China); Meng et al., 2005:27 (ecology, Xin- jiang Uygur Zizhiqu, China); Zhang and Baoping, 2006:109 (behavior and reproductive biology, Xinjiang Uygur Zizhiqu, China). Male. Genitalia. Male D. carinulata can be distinguished from all other species in the D. elongata group by two unique characteristics of the palmate endophallic sclerite (PES): (1) the distal margin is acutely rounded and generally smooth, with one or two small subdistal spines, and (2) the PES has a length to width ratio of 0.61–1.02 (Figs. 17, 32; Table 3). In contrast, the distal margin of the PES is truncate or more broadly rounded with larger distal or subdistal spines in D. elongata, D. carinata, and D. sublineata (Figs. 14–16, 29–31), and the PES is narrowly rounded with a length to width ratio of 0.35–0.63 in D. meridionalis (Figs. 18, 33). The length of the spined area (SL) of the elongate endophallic sclerite (EES) is greater than or equal to 0.33 times (or greater than about one third) the length of the EES in D. carinulata (Figs. 17, 22), compared to the spined area being less than or equal to 0.16 times (or less than about one fifth) the length of the EES in D. elongata (Table 3; Figs. 14, 19, 24, 48). In D. carinulata, the blade of the EES extends less than 2/3 the total length of the sclerite (Figs. 17, 22), and the EES lacks a strong hook at the apex in dorsal aspect (Fig. 27). In contrast, the blade of the EES extends for more than 2/3 the total length of the EES in D. meridionalis (Figs. 18, 23), and the apex of the EES is strongly hooked in dorsal aspect (Fig. 28). Measurements. See Tables 2 and 3. Female. Genitalia. Female D. carinulata can be distinguished from all other members of the D. elongata group by the following combination of characters in the vaginal palpi (VP) and internal sternite VIII (IS VIII): (1) the vaginal palpi are broadly rounded and about as long as wide or longer with a width to length ratio (LP/ WP) of 0.94–1.36 (Fig. 37; Table 4), (2) if the width to length ratio of the vaginal palpi is 0.94, then the width of the widest lobe of the stalk (WLS) of IS VIII is greater than or equal to 0.11 mm (Fig. 37), and (3) the width of the stalk (WST) of IS VIII is 0.36–0.57 mm and this width is 0.49–0.77 times the width of the apical lobe (WAL) (Figs. 37, 42; Table 4). In contrast, the vaginal palpi are triangulate and wider than long with a width to length ratio of 0.46–0.89 in D. carinata and D. sublineata (Figs. 35–36). In D. elongata the vaginal palpi are also broadly rounded but are wider than long with a length to width ratio of 0.52–0.94, and when the width to length ratio of the vaginal palpus is 0.94, the width of the widest lobe of IS VIII is less than or equal to 0.10 mm (Fig. 34). In D. meridionalis, the width of the stalk of IS VIII is narrower than in D. carinulata, measuring 0.22–0.33 mm and this width is from 0.33–0.48 times the width of the apical lobe (Fig. 38, 43). In addition, the width of the widest lobe of the stalk of IS VIII is usually greater than 0.10 mm in D. carinulata (Figs. 37, 42); whereas, in D. meridionalis the width of the widest lobe of the stalk is from 0.04–0.09 mm (Fig. 38, 43; Table 4). Measurements. See Tables 2 and 4. Coloration. Coloration and elytral vittae of dead (Fig. 7) and living (Fig. 8) D. carinulata is very similar to that discussed above for D. sublineata (Figs. 5, 6). Chen (1961) also noted the similar appearance of D. carinulata (as D. e. deserticola) and D. sublineata (as D. e. ab. sublineata) from North Africa. Type material. According to Berti and Rapilly (1973), the Desbrochers type material for D. carinulata, consisting of a male holotype, is deposited in the Demaison Collection at MNHN. As discussed above (see D. elongata – Type material), we were unable to obtain type materials from MNHN. We studied the original description by Desbrochers (1870), which he based on a single individual 5.5 mm in length, the illustrations of the endophallus of the male holotype by Berti and Rapilly (1973), and topotypes from Sarepta, Russia and the neighboring area. Chen’s (1961) type material for D. e. deserticola consists of a holotype male and allotype female from Wei–li (=Yuli), Xinjiang Uygur Zizhiqu, China, and 25 paratypes (8 ♂♂, 17 ♀♀) in Xinjiang from Akesu (= Aksu), Atushen (= Atux), Baicheng, K'u–ch'e (Kuqa), Miquan, Turtiaogou, Shajingzi, Yecheng (= Kargilik), and Wei–li. These are deposited at the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (IZAS). We studied four paratypes (2 ♂♂, 2 ♀♀) from Wei–li County, Xinjiang Uygur Zizhiqu, China (from IZAS), and the original description of D. e. deserticola by Chen (1961).

88 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS We studied the type material of D. koltzei ab. basicornis Laboissière from Khotan (=Hotan), Turkestan (now in Xinjiang Uygur Zizhiqu, China) which consists of a single female according to Laboissière (1935) that Ron Beenen kindly located for us in the Zoological Museum Amsterdam (ZMAN). Material examined. 120♂♂ dissected (diss.), 52♀♀ diss., 199♂♂, 230♀♀, 128 unsexed specimens. AZERBAIJAN: 1♂ diss., Gobustan [Qobustan; 40.08416°N, 49.41583°E], 9-10-VII-2001, V. Dolin, MBPF [2003-07]; 2♀♀ diss., Ordubat [Ordubad; 38.9081°N, 46.0278°E], [19]14, Dr. Veselý, NMPC [2004-26, 2005-37]; 1♀ diss., Ordubad, Araxes [Aras River] Road, Dr. J. Veselý, 9131, NMPC [2004-18]; CHINA: Nei Mongol Zizhiqu (Inner Mongolia): 1♂ diss., Erjina County [specific location not given], 15-VII-1960, Forest Department, Forest Science Institute, Erjina Co., on Tamarix, IZAS [2005-05]; Gansu Sheng: 1♂ diss., 1♀ diss., 1♀, Jiayuguan, 3 km southeast [39.7900°N, 98.3200°E], north side road, 40,238 km odom., 29-VII-1993, C.J. DeLoach, on T. ramosissima, Diorhabda elongata deserticola det. I. Lopatin 2000, 6682–6684, GSWRL [2003-53, 54]; 1♂ diss., 9♀♀, Lanzhou, 4 km east [36.05639°N, 103.84000°E], 39,249 km odom., 26-VII-1993, C.J. DeLoach, on T. hohenackeri [= T. smyrnensis]/T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 6♀♀], 6677–6681, 6729, 8980, GSWRL [2003-48]; Xinjiang Uygur Zizhiqu: 1♂ diss., Bogda Shan Mtns., 80 km northeast Urumqi [Urumchi; 43.81850°N, 87.97160°E]; 4-VI-1993, Mityaev, on saksaul [Haloxylon ammondendron (C.A. Meyer) Bunge (Chenopodiaceae), apparently incidental occurrence], USNM [2003-15]; 1♀ diss., 5♂♂, 8♀♀, Fukang [44.1667°N, 87.9833°E], 8-VIII- 1995 [not 13-VIII-1995 as in some records], Lu Qing-guang, M.T. Liu & Jiang Xi Liang, on T. ramosissima, D. e. deserticola Chen det. I. Lopatin 2000 [1♀], 8584–8586, 8588, 8591–8593, GSWRL(CJD)-1995-25, GSWRL [2005-20] [11 specimens from Lot GSWRL(CJD)-1995-9 identified as prob. D. e. deserticola by R.E. White on 15-XI-1995 at USDA SEL, SEL Lot # 95-10233]; 1♂ diss., 1♂, 2♀♀, Fukang, 8-IX-1996, Li Bao Ping, on T. ramosissima, voucher shipment GSWRL(CJD)-1996-27, GSWRL [2005-90] [9 specimens from Lot GSWRL(CJD)-1996-4 identified as D. elongata by S.M. Clark on 28-X-1996 at West Virginia Dept of Agric.]; 1♂ diss., 1♂, 1♀, Fukang, 8-VI-1996, Li Bao Ping, on Tamarix sp. prob. ramosissima, GSWRL [2006-02]; 1♀ diss., Fukang, southwest, 9-VI-1996, P.E. Boldt, on T. karelinii [T. hispida var. karelinii], GSWRL [2006-01]; 1♂ diss., 11♂♂, 7♀♀, Fukang, Tamarix Park, 1-IX-1997, Li Bao Ping, on Tamarix sp., voucher shipment GSWRL(CJD)-1997-22, GSWRL [2005-100]; 1♂ diss., 1♀ diss., 15♂♂, 10♀♀, Fukang, Tamarisk Park, 18-IV-1998 [not 11-V-1998 as on some shipment records and labels], Li Bao Ping, on T. ramosissima, 805, 806, 808, 811, 812, 814–815, 820, 821, 823–825, voucher shipment GSWRL-1998-05, GSWRL [2005-67, 68] [parasite Erynniopsis antennata Rondani (Diptera: ) emerged from 21 adult specimens in Lot GSWRL-2000-04 identified by N. E. Woodley on 23-II-2000 at USDA SEL, SEL Lot # 0001060]; 1♂ diss., 4♂♂, 8♀♀, Fukang, 13-VII-1999, Li Bao Ping, on Tamarix spp., voucher shipment BCW/WRRC-99-04 received 21-VII-1999, GSWRL [2005-75] [7 specimens from Lot GSWRL-1999-2 identified as D. e. deserticola by S. Lingafelter on 23-VII-1999 at USDA SEL, SEL Lot # 9904542] [USDA lab colony at Albany, CA] [introduced into cages in Nevada, Colorado, and Wyoming in 1999; released in 2001] [used in biological studies of DeLoach et al. (2003b), Lewis et al. (2003a, 2003b), Petroski (2003), Cossé et al. (2005), Herrera et al. (2005), Milbrath and DeLoach (2006a), Herr et al. (in prep.), Bean and Keller (in prep.), and Thompson et al. (in prep.)] [in field studies by DeLoach et al. (2004), Dudley and Kazmer (2005), Aber et al. (2005), USDA-APHIS (2005), and Dudley et al. (in prep.)]; 1♂ diss., 2♀♀ diss.; Fukang, 13-VII-1999, Li Bao Ping, on Tamarix spp., voucher shipment BCW/WRRC-99-04, USDA/ARS lab colony at Temple, Texas (ex Lovell, WY, 2002, J.L. Tracy), GSWRL [2003-28, 29, 32]; 1♂ diss., 1♂, 3♀♀, Fukang, ca. 3 km west [44.14240°N, 87.92806°E], km marker 748, 15-VII-1992, R.Wang & C.J. DeLoach, on T. arceuthoides [1♂ diss], on T. hispida var. karelinii – no flowers [1♂, 2♀♀], on T. laxa – no flowers [1♀], D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 2♀♀], 8974, GSWRL [2005-18]; 2♀♀ diss., Fukang, ca. 3 km west, km marker 749, 15-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima very few flowers [1♀ diss.], on T. ramosissima foliage [1♀ diss.], D. e. deserticola det. I. Lopatin 2000, 6728, 6659, GSWRL [2005-12,13]; 1♂ diss., 3♂♂, 2♀♀, Fukang, 7 km west [44.13658°N, 87.87737°E], 10,406 km odom., 7-VII- 1992, R. Wang & C.J. DeLoach, on T. ramosissima [1♂ diss., 2♂♂, 1♀], on T. laxa - no flowers [1♀], D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♂♂, 1♀], 6646–6649, GSWRL [2005-17]; 4♂♂ diss., 6♂♂, 6♀♀,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 89 Fukang, 18 km west [44.11252°N, 87.77169°E], 14,731 km odom., 9-VIII-1993, C.J. DeLoach, on T. leptostachya [1♂ diss., 2♂♂, 1♀], on T. karelinii [hispida var. karelinii]-some flowers (red) [1♂ diss., 1♂, 2♀♀], on T. ramosissima-some flowers [1♂ diss., 1♂, 1♀], on T. elongata [1♂ diss., 1♂, 1♀], T. laxa – no flowers [1♂, 1♀], D. e. deserticola det. I. Lopatin 2000 [4♂♂ diss., 3♂♂, 4♀♀], 6662–6669, 6693, 6694, GSWRL [2003-47, 2005-07, 19, 22]; 1♂ diss., Jiashi [39.4883°N, 76.7125°E], 11-VII-1992, R.Wang & C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000, 6515, GSWRL [2005-04]; 1♀ diss., Jiashi, 9 km west [39.45198°N, 76.61615°E], 11-VII-1992, R. Wang and C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000, 6658, GSWRL [2005-05]; 1♂ diss., Khotan [Hotan; 37.09972°N, 79.92694°E], D. elongata ab. sublineata Lucas, HNHM [2003-14]; 1♂ diss., 1♂, 1♀, Khotan [Hotan], Kashgar, 13-VI-1890, B. Grombchevskii, ZIN [2005-02]; 1♀, Khotan [Hotan], 6-12-V-1930, J.A. Sillem, Diorhabda koltzei Weise ab. basicornis Laboissière, TYPE [Laboissière 1935], Nederlandsche Karakorum Expeditie, ZMAN [2008-01]; 1♂ diss., 2♂♂, 4♀♀, Kuldsha [Yining; 43.9000°N, 81.35000°E], Daharkent, NHMB [2003-27]; 1♂ diss., Sanchakou, 50 km west [39.93778°N, 77.8600°E], 10-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima, D. e. deserticola det I. Lopatin 2000, 6657, GSWRL [2003-57]; 2♂♂ diss., 1♂, 4♀♀, Shanshan [42.8667°N, 90.1667°E], edge of, 41,571 km odom., 5-VIII-1993, C.J. DeLoach, on T. laxa, D. e. deserticola det I. Lopatin 2000, 6671, 6673, 6674, 6676, 6689, 6690, 6730, GSWRL [2005-10, 96]; 1♂ diss., 4♂♂, 1♀, Turpan [Turpan Botany Station; 10 km SE Turpan; 42.863°N, 89.222°E], 17-VII-1992, C.J. DeLoach, on T. arceuthoides, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂], 8975, 8978, voucher shipment GSWRL(CJD)-1992-21, GSWRL [2005-15] [12 specimens from Lot GSWRL(CJD)-1993-11 identified as prob. D. e. deserticola by R.E. White on 15-X-1993 at USDA SEL, SEL Lot # 93-10349]; 1♂ diss., 2♂♂, 1♀, Turpan Botany Station [10 km SE Turpan], 17-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 1♀,], 6660–6662, voucher shipment GSWRL(CJD)-1992-21, GSWRL [2005-16] [used in biological studies of DeLoach et al. (2003b)]; 3♂♂ diss., 2♀♀ diss., Turpan Botanical Garden, 10 km SE Turpan, 19-VI-2002, Li Bao Ping, on Tamarix spp., voucher USDA/ARS lab colony at Temple, Texas (4-IX-2002, J.L. Tracy; 1♂, 1♀, 25-26-VIII-2003, L. Milbrath [nos. 1265, 1268]), shipment EIWRU-2002-1005, GSWRL [2003-21, 31, 33, 2004-04, 2005-84] [released in Colorado in 2004] [used in biological studies of Milbrath and DeLoach (2006a, 2006b), Herr et al. (in prep.), and Bean and Keller (in prep.)]; 1♂, Turpan, ca. 30 km east, 41,751 k m odom., 5-VIII-1993, C. J. DeLoach, 20 sweeps on T. laxa, GSWRL ; 1♀, Turpan, 74 km northwest [43.20415°N, 88.46102°E], White Poplar Valley, rd. km 687, 7-VIII-1993, C.J. DeLoach, on T. arceuthoides, D. e. deserticola det. I. Lopatin 2000, 6691, GSWRL; 1♂ diss., 1♂, 4♀♀, Turpan, 76 to 164 km northwest, 7-VIII-1993, C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 1♂, 3♀♀], 8981–8984, voucher shipment GSWRL(CJD)-1993-15, GSWRL [2005-76] [11 specimens from Lot GSWRL(CJD)-1993-11 identified as prob. D. e. deserticola by R.E. White on 15-X-1993 at USDA SEL, SEL Lot # 93-10349] [used in biological studies of DeLoach et al. (2003b)]; 1♂ diss., 5♂♂, 9♀♀, Turpan, 74 to 164 km northwest, White Poplar Valley, 7-VIII-1993, C.J. DeLoach, on T. arceuthoides and T. ramosissima, GSWRL [2004-14]; 1♂ diss., Urumqi [43.8000°N, 87.58333°E], water reservoir in suburb, VII-1998, Li Bao Ping, on Tamarix sp., voucher shipment GSWRL-1998-12, received 4-VIII-1998, GSWRL [2005-24]; 1♂ diss., 2♂♂, 2♀♀, Urumqi, 12 km northeast [43.89406°N, 87.61073°E], by fertilizer factory, 42,403 km odom., 8-VIII-1993, C.J. DeLoach, T. hispida, D. e. deserticola I. Lopatin 2000 [1♂ diss., 2♂♂, 1♀], 6685–6688, GSWRL [2005-01]; 1♂ diss., 1♀, Urumqi, 47 km southeast [43.54156°N, 87.96419°E], 10,525 km odom., 8-VII-1992, R.Wang & C.J. DeLoach, on T. ramosissima - flowers, D. e. deserticola det. I. Lopatin 2000, 8973, GSWRL [2005-11]; 1♂ diss., 2♂♂, 4♀♀, Urumqi, 57 km east [43.56707°N, 88.07503°E], 8-VII-1992, R. Wang & C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♂♂, 3♀♀], 6645, 6650, 6651, 6653–6656, GSWRL [2005-06]; 1♂ diss., Urumqi, 47 km southwest [43.44176°N, 87.23066°E], 10,525 km odom., 8-VII- 1992, C.J. DeLoach, on T. ramosissima, GSWRL [2005-37]; 1♀ diss., Wei Lei Co. Farm 6 [near Yuli: 41.33060°N, 87.25780°E], 14-V-1960, Wang Shuyong, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-04]; 1♂ diss., Wei Lei Co. Farm 6 [near Yuli], 14-V-1960, Zhang Facai, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-03]; 1♀ diss., Wei Lei Co. Farm 4 [near Yuli], 17-V-1960, Wang Shuyong, D. e.

90 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS deserticola det. Chen, PARATYPE, IZAS [2005-02]; 1♂ diss., Wei Lei Co. Farm 4 [near Yuli], 17-V-1960, Zhang Facai, D. e. deserticola det. Chen, PARATYPE, IZAS [2005-01]; IRAN: 3♂♂ diss., 4♀♀ diss., 1♂, 3♀♀, 124 specimens [NMPC], Abaregh [Abareq; 29.31667°N, 57.91667°E; irrigation channel in semi-desert; T. leptopetela (= T. kotschyi) present (Hoberlandt 1981)], 53 km northwest Bam, 25-IV-1973, Exp. Nat. Mus. Praha Loc. No. 178, NMPC [2004-30, 31, 40, 2005-04, 05], USNM [2003-06, 2004-04]; 1♂ diss., 1♀ diss., Bareng [Berang] to Nasratabad [Nosratabad] [app. location: 30.23670°N, 61.00170°E], Sistan, 11-V-1898, N. Zarudnyl, ZIN [2004-18, 21]; 1♂ diss., Bendun [Bandan] [to] Nayzar [Naizar marsh] [app. location: 31.3033°N, 60.8292°E], Sistan, east Persia, 8-V-1898, N. Zarudnyl, 17585, ZIN [2005-05]; 2♂♂ diss., 5♂♂, Kashan, 25 km toward Natanz [Natenz] [33.81444°N, 51.74361°E], 3,200 m elev. 2-V-2000, J. Gaskin, on Tamarix sp., #772, GSWRL [2003-06, 37]; 1♂ diss., 1♀, Khatunabad [29.46667°N, 57.80000°E; stony semi- desert; Tamarix sp. not recorded (Hoberlandt 1981)], 70 km northwest Bam, 25-IV-1973, Exp. Nat. Mus. Praha Loc. No. 179, NMPC [2004-35b]; 1♂ diss., Khorassan [Ostan-e Khorasan Province; on northeast border of Iran; specific locality not given], 12-13-VIII-1901, N Zarudnyl, ZIN [2004-01]; 1♂ diss., 1♀, Minu Dasht, 60 km east [37.39°N, 55.85°E], MRSW Park [Mohammed Reza Shah Wildlife Park (= Golestan Biosphere Reserve)], 28-VIII-1972, [on T. ramosissima (Gerling and Kugler 1973)], D. elongata det. S.L. Shute 1973, P27-29, TAUI [2003-03]; 1♂ diss., 1♀ diss., Shakhrud K. [Sharud Kola; 36.41670°N, 52.85000°E], north of Gorgan [southeast of Astrabad], Christof, ZIN [2004-17, 20]; KAZAKHSTAN: 1♂ diss., 2♂♂, 2♀♀, Chilik [Shelek], near [2♂♂, 2♀♀], or 25 km east [43.6715°N, 78.5050°E], Buryndysu environs [1♂ diss.], 6-VI-1999, Mityaev & Jashenko, on T. ramosissima, voucher shipment GSWRL-1999-8, GSWRL [2004-03] [introduced near Delta, Utah in 1999] [28 specimens from Lot GSWRL-1999-1 identified as D. elongata by A. Konstantinov on 20-VII-1999 at USDA Systematic Entomology Laboratory (SEL), SEL Lot # 9904498]; 1♂ diss., 4♂♂, 5♀♀, Bzandy-Su [Buryndysu] Campsite, 3-4 km north [43.70650°N, 78.68490°E], 186 km east northeast Almaty, 99,921 km odom., 4-VI-1995, C.J. DeLoach, on T. ramosissima, D. e. deserticola det I. Lopatin 2000 [2♂♂, 1♀], 8989–8891, GSWRL [2003-56]; 1♂ diss., 9♂♂, 6♀♀, Chilik [Shelek; 43.60030°N, 78.29630°E], near, 106 east of Almaty, 8-VIII-1995, I.D. Mityaev, on T. ramosissima, D. elongata det. I.D. Mityaev [1995; 3♂♂, 4♀♀], voucher shipment GSWRL(CJD)-1995-23, GSWRL [2005-91]; 1♂ diss., Chilik, near, 4-VI-1997, I.D. Mityaev and P.E. Boldt, on T. sp. prob. ramosissima, voucher shipment GSWRL(CJD)-1997-13, GSWRL [2005-92]; 1♂ diss., 5♂♂, 8♀♀, Chilik [Shelek], near, 1-VIII-1999, I. Mityaev & R. Jashenko, on T. ramosissima, voucher shipment GSWRL-1999- 15, GSWRL [2005-98]; 1♂ diss., 3♂♂, 1♀, Chilik [Shelek], environs, 21-V-2000, Mityaev & Jashenko, on T. ramosissima, voucher shipment GSWRL-2000-9, GSWRL [2005-80, 97] [used in biological studies by DeLoach et al. 2003b and Lewis et al. (2003a)]; 2♂♂ diss., 8♂♂, 8♀♀, Chilik [Shelek], 7 km before [southwest; 43.49000°N, 78.25000°E], 130 km east Almaty, site 1, 99,865 km odom., 3-VI-1995, C.J. DeLoach, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [6♂♂, 2♀♀], 8595–8664 , 8985, 8987, 8988, voucher shipment GSWRL(CJD)-1995-11, GSWRL [2003-45, 2005-14] [10 specimens from Lot GSWRL(CJD)-1995-9 identified as prob. D. e. deserticola by R.E. White on 15-XI-1995 at USDA SEL, SEL Lot # 95-10233]; 2♂♂ diss., 2♂♂, 5♀♀, Chilik [Shelek], 38 km east [Buryndysu; 43.69710°N, 78.66300°E], 15-VI-1996, P.E. Boldt, on T. ramosissima, D. e. deserticola det. I. Lopatin 2000 [1♂ diss., 2♀♀], 9013 [1♂ diss.], 9015, 9016, voucher shipment GSWRL(PEB)-1996-9, GSWRL [2003-83, 2005-23]; 1♂ diss., Chilik [Shelek], 23 km east [odom. reading 9,465 km] and 38 km east [odom. reading 9,480 km; Buryndysu; not west Chilik as in some shipment records], 15-VI-1996, I.D. Mityaev & P.E. Boldt, on T. ramosissima, voucher shipment GSWRL(PEB)-1996-9, GSWRL [2005-99] [8 specimens from Lot GSWRL(CJD)-1996-4 identified as D. elongata by S.M. Clark on 28-X-1996 at West Virginia Dept of Agric.] [8 specimens (no. 9205) from Lot GSWRL(CJD)-1997-1 identified as D. e. prob. elongata by A. Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023]; 2♂♂ diss., 5♀♀ diss., 3♂♂, 3♀♀, Chundzha, 40 km west [43.47035°N, 79.02773°E], desert near Boguty Mtns., 9-VI-1999, Mityaev & Jashenko, on T. ramosissima, voucher shipment GSWRL-1999-9, GSWRL [2003-62, 63, 64, 2005-02, 48, 73, 74] [subject of field studies by Mityaev and Jashenko (1999, 2007)]; 2♂♂ diss., 3♂♂, 3♀♀, Chundzha, 40 km west, desert near Boguty Mtns., 1-VIII-1999, I.D. Mityaev & R. Jashenko, on T. ramosissima, voucher shipment GSWRL-1999-15,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 91 GSWRL [2005-93, 95]; 1♂ diss., Dzhulek [Zhulek; 44.28330°N, 66.43330°E], Syrdar’indkaya Oblast, VII- 1910, Kozchanchikov, ZIN [2004-15]; 1♂ diss., Indersk [Ynderbor; 48.55000°N, 51.78330°E], Reitter, D. e. ab. sublineata, HNHM [2003-10]; 7♂♂ diss., 2♂♂, 1♀, Kerbulak [43.9311°N, 77.6019°E], ca. 150 km north northeast Almaty, Ili River Valley, 2-VI-1999, Mityaev & Jashenko, on Tamarix sp., voucher shipment GSWRL-1999-7, GSWRL [2004-19; 2008-06, 07, 08, 09 ,10, 11] [subject of field studies by Mityaev and Jashenko (1999, 2007)]; 1♂ diss., Lavar [43.56860°N, 78.13360°E], 10 km west Chilik [Shelek], 21-V-2000, Mityaev & Jashenko, on T. ramosissima, voucher shipment GSWRL-2000-9, GSWRL [2004-21] [introduced near Delta, Utah in 2000]; 1♂ diss., 10♂♂, 3♀♀, Masaq [43.62028°N, 78.30556°E] [not Marak as in some shipment records and labels], near, Ili Desert, 4-VI-1997, I.D. Mityaev and P.E. Boldt, on Tamarix sp., voucher shipment GSWRL(CJD)-1997-13 [1♂], GSWRL [2004-18]; 1♂ diss., Sarytogay, Charyn Canyon [43.43240°N, 79.15040°E], 13-VI-1998, Mityaev, on Tamarix sp., voucher shipment GSWRL-1998-10, GSWRL [2004-20]; 1♂ diss., Shelek, near, Shelek River, 6-VI-1999, Mityaev & Jashenko, on T. ramosissima, voucher shipment GSWRL-1999-8, GSWRL [2004-02] [introduced near Delta, Utah in 1999] [used in biological studies of DeLoach et al. (2003b) and Lewis et al. (2003b); subject of field studies by Mityaev and Jashenko (1999, 2007)]; KYRGYZSTAN: 1♂ diss., 1♀ diss., Dostuk [41.36972°N, 75.63667°E], environs, At–Bash River bed, Naryn Region, 20-VIII-1995, S. Saluk, USNM [2003-14, 16]; 1♂ diss., Naryn, 80 km west [41.36972°N, 75.3799°E], Central Tian Shan Mts., 3-VII-1966, E. Gurieva, ZIN [2004-14]; MONGOLIA: 1♂ diss., Mongolia [specific locality not given], Potanin, Coll[ection] Weise, [D.] e. var. sublineata [1♂ diss.], ZMHB [2006-05] [probably from series collected by G.N. Potanin from 22-VII to 3- VIII-1886 from Central Mongolia listed as D. e. var. sublineata by Weise (1890)]; 1♂ diss. 1♀ diss., southwestern Mongolia [specific locality not given], 17-VI [1♂ diss.], 18-VI [1♀ diss.] [circa 1928 (Hedin 1943)], Söderbom, D. elongata [1♂ diss.], Sven Hedins Expedition Central Asia, NHRS [2003-02, 03]; Bayanhongor Aymag: 1♂ diss., Tooroyn Bulag spring [42.76745°N, 98.96116°E], 13 km east Tsagan Bulag, Bayanhongor Aymag, 16-VIII-1969, Zaitsev, ZIN [2004-16]; Hovd Aymag: 1♂ diss., Baruun Hurray [Barun Khurai Tract; 45.66667°N, 91.66667°E], 50 km southwest of Uench [Uyonch], 3-4-VIII-1976, L.N. Medvedev, D. elongata det. L. Medvedev, MRSN [2003-01] [from series listed as D. elongata by Medevedev and Voronova (1979)]; 2♂♂ diss., 2♀♀, Bulgan Somon [Burenhayrhan; 45.99394°N, 91.55267°E], 10 km south southwest, Chovd Aimak [Hovd Aymag], 4-5-VII-1966, Exp. Dr. Z. Kaszab, Nr. 628, D. elongata sublineata det. K. Lopatin [1♂ diss., EGRC], USNM [2003-08], EGRC [2005-05] [from series listed as D. e. sublineata by Lopatin (1968)]; 1♂ diss., Yelkhon, 20 km southeast [sic; should be southwest along Bodochiin Gol from Altai] Altai [Bor–Udzuur], on Bodonchiyn [Bodochiin Gol stream] [45.7008°N, 92.0836°E], 27- VII-1970, Emeljanov, ZIN [2004-05]; RUSSIA: 1♂ diss., 1♂, 1♀, Astrachan [Astrakhan; 46.34944°N, 48.04917°E], coll[ection] Koltze, 10883, DEI [2003-25]; 1♀ diss., signata [designated] Astrachan, Becker, coll[ection] L.V. Heyden, [Galeruca] carinulata Desbr., DEI [2003-04]; 1♀ diss., 2♂♂, 1♀, Astrachan, Coll[ection] Kraatz, [Galeruca] carinulata Desbr. [1♀ diss.], DEI [2005-11]; 1♂, 1♀, Astrakan [Astrakhan], Becker, D. e. ab. carinata det. Le Moult, IRSNB; 1♂ diss., 1♀, Astrakhan, Letzner, coll[ection], Gal[eruca] carinulata Desbr.[1♂ diss.], DEI [2003-03]; 1♂ diss., 1♂, 2♀♀, Astrakhan, Becker, coll[ection] Reitter, D. elongata, HNHM [2003-01]; 1♂ diss., 1♀, Astrakhan. [region; specific locality not given], Bekker, ZIN [2005-04];1♂ diss., Derbent [42.06278°N, 48.29583°E], Coll[ection] Koltze, 10884, DEI [2003-05]; 1♀ diss., Derbent, Becker, DEI [2004-01]; 1♂ diss., 1♂, 1♀, Malaya Areshkeva [44.02083°N, 46.82305°E], northern Daghestan [Respublika Dagestan], 31-V-1925, A.N. Kiritschenko, ZIN [2004-03]; 1♂ diss., 1♀ diss., Sarepta [48.52778°N, 44.48361°E], G. carinulata Desbrochers [1♂ diss.], Coll[ection] Stierlin, DEI [2003-07, 26]; 1♀ diss., Sarepta, Becker, [G. ] carinulata, NHRS [2003-01]; 1♀ diss., Sarepta, D. Becker, ex. coll. J. Weise, ZMHB [2003-01]; 1♀ diss., Sarepta. Schioda [?], 896, Collectie C. et O. Vogt Acq. 1960, ZMAN [2008-13]; 2♂♂ diss., 1 ♀ diss., Ikrianoe [Ikryanoye; 46.09440°N, 47.73470°E] Village, Astrakhan region, 22–28-VIII- 2002, Osipov collection, GSWRL [2003-70, 2004-25, 26]; TAJIKISTAN: 1♂ diss., 1♀ diss., Vakhsh River, Tigrovaya Balka State Reserve [37.10528°N, 68.30389°E], Dzhilikul District, Kurgan–Tube [Kurgan Tyube] Region, 9-V-1992, Osipov collection, GSWRL [2003-77, 2004-08]; TURKMENISTAN: 2♂♂ diss., 2♀♀ diss., Ashgabat, 2-IX-2002, S.N. Myartseva, on T. ramosissima, GSWRL [2003-01, 03, 05, 2006-06]; 1♀

92 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS diss., Ashgabat, Dry Sport Lake, 11-V-1997, S. N. Myartseva, on Tamarix ramosissima [prob. T. aralensis J. Gaskin], voucher shipment GSWRL(CJD)-1997-11, GSWRL [2005-101] [8 specimens (nos. 9202, 9203) from Lot GSWRL(CJD)-1997-1 identified as D. e. prob. elongata by A. Konstantinov on 21-V-1997 at USDA SEL, SEL Lot # 9704023]; 2♀♀ diss., Ashgabat [37.95°N, 58.36667°E], Dry Sport Lake, 11-VI-1997, S. N. Myartseva and P. Boldt, on T. ramosissima [prob. T. aralensis J. Gaskin], voucher shipment GSWRL(CJD)- 1997-12, D. elongata det. I. Lopatin 1999 [1♀], 9023, GSWRL [2005-08, 2006-07]; 2♂♂ diss., Ashgabat, Dry Sport Lake, 26-IX-2002, S.N. Myartseva, on Tamarix spp., GSWRL [2006-08, 09]; Ashgabat, near Quaragum Canal, 27-V-1994, C.J. DeLoach, on Tamarix sp., D. e. deserticola det. I. Lopatin 2000, 6731, GSWRL [2005-03]; 1♀ diss., 2♂♂, Ashgabat, canal [Quaragum], 11-VI-1997, S.N. Myartseva and P. Boldt, on Tamarix sp., D. elongata det. I. Lopatin 1999, 9017–9019, GSWRL [2005-21]; 3♂♂ diss., 4♀♀ diss., Ashgabat, Karakum [Quaragum] Canal, 12-IX-2002, S. N. Myartseva, on T. ramosissima, GSWRL [2003- 13b, A5, A6, A7, A8, 2005-49, 63]; UNKNOWN COUNTRY: 1♀ diss., Araxesthal [Aras River of Armenia, Azerbaijan and Iran], Reitter, HNHM [2003-11]; 1♂ diss., Caucasus [specific locality not given; possibly Armenia or Azerbaijan], Reitter. Leder., NMPC [2005-41]; 1♀ diss., 1♀, Kammenaja retschka [Kamennaya Rechka, Russia?], 4-VI-1909, Collectie C. et. O. Vogt Acq. 1960, ZMAN [2008-03]; 1♀ diss., Tscholokai [Tschkalow/Orenburg, Russia?], 31-V-1909, Collectie C. et. O. Vogt Acq. 1960, ZMAN [2008-02]; UNITED STATES OF AMERICA (introduced): Colorado: Pueblo Co.: 1♂ diss., 4♂♂, 7♀♀, Arkansas River, Pueblo Reservoir, 38.270°N, -104.713°W, 7-IX-2005, D. Hosler and F. Nibling, on T. ramosissima, voucher [source: Fukang, China], GSWRL [2006-16]; Nevada: Churchill Co.: 1♂ diss., 5♂♂, 4♀♀, Carson Sink, Trinity Junction, 38.92°N, -118.75°W, 23-IX-2005, T. Dudley, on T. ramosissima, voucher [source: Fukang, China], GSWRL [2006-12]; Mineral Co.: 1♂ diss., 4♂♂, 5♀♀, Walker Lake, 38.88°N, -118.79°W, 21-IX-2005, T. Dudley, on T. ramosissima, voucher [source: Fukang, China], GSWRL [2006-13]; Pershing Co.: 1♂ diss., 3♂♂, 6♀♀, Humboldt Sink, 40.13°N, -118.42°W, 23-IX-2005, T. Dudley, on T. ramosissima, voucher [source: Fukang, China], GSWRL [2006-14]; Utah: Millard Co.: 1♂ diss., 5♂♂, 6♀♀, Sevier River west of Delta, 39.172°N, -112.903°W, 15-IX-2005, C.J. DeLoach, on T. ramosissima, voucher [source: Shelek, Lavar, and environs Buryndysu, Kazakhstan], GSWRL [2006-17]; Wyoming: Big Horn Co.: 1♂ diss., 4♂♂, 5♀♀, Big Horn Canyon National Recreation Area [44.857°N, -108.208°W, east of Lovell], 22-VIII-2005, D. Kazmer, on T. ramosissima, voucher [source: Fukang, China], GSWRL [2006-15]; UZBEKISTAN: 2♂♂ diss., 11♂♂, 2♀♀, Buchara [Buxoro], southeast [39.7555°N, 64.4867°E], 3-IV-1999, R. Sobhian, on Tamarix spp., D. elongata det. I.K. Lopatin 1999 [12♂♂, 2♀♀], 9036–9048, 9050, GSWRL [2003-82, 2008-20]; 2♂♂ diss., 1♀ diss., Buxoro, 5 km northwest [39.8228°N, 64.39134°E], along the road to Gazli, 27-IX-2002, R. Sobhian, on Tamarix, USDA lab [Buchara] colony at Albany, California, voucher (2002–2003, D. Bean), shipment EIWRU-2002-1010, GSWRL [2005-53, 54, 56]; 2♂♂ diss., 1♀ diss., 5♂♂, 2♀♀, Buchara [Buxoro], 10 km west [39.7614°N, 64.3011°E], 3-IV-1999, Kirk & Sobhian, on Tamarix sp., D. elongata det. I.K. Lopatin 1999 [2♂♂ diss. 4♂♂], 9028–9031, 9033, 9034, GSWRL [2003-80, 2004-05, 2008-21]; 1♂ diss., 1♂, Bukhara [Buxoro], 25 km west [39.69778°N, 64.18315°E], 20-V-1994, Osipov Coll., GSWRL [2004-22]; 1♂ diss., 1♂, Karshi [Qarshi], 10 km west [38.92278°N, 65.73307°E], 7-IV-1999, A. Kirk & R. Sobhian, on Tamarix, D. elongata det. I.K. Lopatin 1999, 9065, 9066, GSWRL [2003-78]; 1♂ diss., Tashkent [Toshkent], 140 km south [approximate location: 40.29057°N, 68.91332°E], 10-IV-1999, Kirk & Sobhian, on Tamarix spp., GSWRL [2004-17]. Distribution. General. The native distribution of D. carinulata ranges from Sarepta, Russia south to Iran and east to China and Mongolia (Map 5). Surveys by our colleague C. J. Deloach reveal that D. carinulata populations are temporally sporadic in China and Kazakhstan, with outbreaks in some years and rarity in other years. Further collections might also reveal D. carinulata in Armenia, Afghanistan, Pakistan and northwestern Iraq. Confirmed Records. We dissected specimens (see Material examined) from Russia, the single country with a previous literature record for D. carinulata (Desbrochers 1870) (Map 5). New Records. We have dissected specimens from the following countries for which we find no previous specific reports of D. carinulata in the literature (Maps 5–6): Azerbaijan, Iran, Turkmenistan, Kazakhstan,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 93 Uzbekistan, Tajikistan, Kyrgyzstan, China, Mongolia, and the United States of America (Nevada, Utah, Colorado, Wyoming; introduced; Map 7, see Potential in Tamarisk Biological Control below for additional details). Past reports of D. elongata from Azerbaijan, Turkmenistan, Kazakhstan, Uzbekistan, and the United States should refer, at least in part, to D. carinulata. Past reports of D. e. deserticola from China, Mongolia, Kazakhstan, and the United States should also refer to D. carinulata (see synonymy above). Past reports of D. e. sublineata and D. e. ab. sublineata in China and Mongolia are also D. carinulata. Unconfirmed Records. We find no country distribution records that remain unconfirmed. We consider reports of Diorhabda from Littlefield, Arizona (L.D. Walker, USDI Bureau of Land Management, St. George, Utah) that spread south from Utah as D. carinulata. All available locations from which specimens were dissected from China (25) and Mongolia (6) were D. carinulata, with the exception of specimens of D. carinata from China near the border of Kazkhstan in the Ili Kazakh Autonomous Prefecture and Yining. Therefore, we consider all literature records of D. elongata, D. e. sublineata and D. e. deserticola from these countries to be D. carinulata, although D. carinata may also be present at sites in far western Xinjiang Uygur Zizhiqu, China. From 11 locations east of 75° E in Kazakhstan, D. carinulata was dissected from ten locations; while, D. carinata was dissected from four locations, two of which were series greatly predominated by D. carinulata. Therefore, we consider all literature reports of D. elongata east of 75°E in Kazakhstan to include D. carinulata, but it is probable that D. carinata occurs in lower frequency at many of these same sites. Only D. carinulata was dissected from all five locations available from north of 42°N in southern Russia and we consider all reports of D. elongata from this area to be D. carinulata. Below are 33 unconfirmed locality records that we consider as D. carinulata, but are listed as D. elongata in Russia, Kazakhstan and Mongolia, D. e. sublineata in Mongolia (Lopatin 1970, 1977b), and D. e. deserticola in China (Map 5): CHINA: Gansu Sheng: Anxi [Yuanquan; 40.500°N, 95.800°E], on Tamarix; Dunhuang [40.16667°N, 94.68333°E], on Tamarix; Nei Mongol Zhizhiqu: Dong He River [41.73647°N, 100.85793°E], on Tamarix (Bao Ping, Nanjing Agricultural University, Nanjing, China, pers. comm.); Ningxia Huizu Zizhiqu: Yanchi [37.7869°N, 107.3994°E], on Tamarix (B. Ping, pers. comm.); Xinjiang Uygur Zizhiqu: Akesu [Aksu; 41.1231°N, 80.2644°E]; Atushen [Atux; 36.7061°N, 76.1519°E]; Baicheng [41.7739°N, 81.8689°E] (Chen 1961); Hami [42.800°N, 93.45000°E] (Sha and Yibulayin 1993); Hotan (Sha and Yibulayin 1993); K'u–ch'e [Kuqa; 41.7278°N, 82.93640°E]; Miquan [43.96667°N, 87.7000°E]; Shajingzi [40.76667°N, 79.96667°E]; Turtiaogou [geocoordinates not locatable] (Chen 1961); Turpan [D. carinulata dissected from same location], on Tamarix (Sha and Yibulayin 1993); Yecheng [Kargilik; 37.88500°N, 77.4131°E]; Wei–li [Yuli; 41.3306°N, 86.25780°E] (Chen 1961); KAZAKHSTAN: Bakanas [Baqanas; 44.8081°N, 76.2772°E], Ili River, on Tamarix (Ishkov 1996); Boguty [desert near Boguty Mts., 40 km west of Chundzha; D. carinulata dissected from same location], on Tamarix (Mityaev and Jashenko 1999, 2007); Borokhudzhir [43.9667°N, 79.5833°E], on Tamarix (Kulenova 1968); Burundysu [Chilik, 38 km east; D. carinulata dissected from same location], on Tamarix; Chilik [Shelek; D. carinulata dissected from same location], near, in Chilik River Valley, on Tamarix; Kerbulak [D. carinulata dissected from same location], Ili River Valley, on Tamarix (Mityaev and Jashenko 1999, 2007); Koksu [45.02780°N, 77.9460°E], on Tamarix (Kulenova 1968); Lavar [D. carinulata dissected from same location], on Tamarix (Mityaev and Jashenko 1999, 2007); Masak [Masaq; D. carinulata dissected from same location] Town environs, Chilik [Shelek] riverbed, 22-VI-1995, Myricaria sp. (Mityaev and Jashenko 1997, 2007); Sarytogay [D. carinulata dissected from same location], Charyn Canyon, on Tamarix (Mityaev and Jashenko 1999, 2007) ; MONGOLIA: Bayanhongor Aymag: 2 specimens, Echin Gol Oasis [Ekhiin Gol, 43.30851°N, 98.99646°E], 90 km north of Capanbulag border post, 950 m, 27–28-VI- 1967 (Nr. 855), on Tamarix (Lopatin 1970); Ezhiin Gola [Ekhiin Gol], 900 m, 3–4-IX-1976; Khatan–Sudlyn–Bulak [geocoordinates not locatable], 75 km west north west Tsagan–Bulak [Tsagaan Bulag spring], 1,450 m [elev.], 28-VII-1977 (Medvedev and Voronova 1979); Zxiln Gol oasis [geocoordinates not locatable], 150 km south Shine Dzhinsta [Dzalaa or Shinejinst], 900 m [elev.], 12–15-IX-1976 (Medvedev and Voronova 1977b); Talyn–Belgekh–Bulak spring [geocoordinates not locatable], 40 km south southeast of Ezhiin Gola [Ekhiin Gol], 1,250 m [elev.], 26-VII-1977 (Medvedev and Voronova 1979); Govi–Altay

94 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Aymag: Dzahuy [44.950°N, 96.600°E]; Khatan Khaïrkhan environs [Hatan Hayrhan Mt.], 50 km east southeast Altay [geocoordinates not locatable] (Medvedev and Voronova 1977b); Khatan Khayrkhav [Hatan Hayrhan Mt.], north slope, 1,225 m [elev.] [44.9881°N, 96.1792°E], 13–14-VII-1976; Khatan Khayrkhav [Hatan Hayrhan Mt.], 20 km southwest, 1,300 m [elev.] [44.8505°N, 96.0897°E], 15-VII-1976 (Medvedev and Voronova 1977a); Shara–Xulsni–Bulak spring [Shara Hulasanii Bulag; 43.3°N, 97.75°E], 100 km west Ezhiin Gola [Ekhiin Gol], 29-VII-1977; Hovd Aymag: Bodonchiyn Gol [Bodochiin Gol stream], 15 km southeast [sic; should be southwest along Bodochiin Gol from Altai] Altai [Bor–Udzuur] [45.7367°N, 92.1217°E], 4-VII-1976, on Myricaria alopecuroides (Medvedev and Voronova 1979); Jarantaj [geocoordinates not locatable], Wuste, 15-V-1974, R. Piechocki (Lopatin 1977b); Uench Gol [Uyonch Gol stream], 25 km southwest Uencha [Uyonch], 1,300 m [elev.] [45.8861°N, 91.7807°E], 28-VII-1976 (Medvedev and Voronova 1977a); Omnogovi Aymag: Obooto Hural, 50 km west [43.02078°N, 100.90040°E], 1600 m [elev.], 25-VII-1977 (Medvedev and Voronova 1979); RUSSIA: Dosang [46.9044°N, 47.9111°E], Astrakhan Province, VI-2003, K.A. Grebennikov (with photograph, Berlov 2006). From our habitat suitability index models (Maps 8b and 9a) and our preliminary species distribution models based on 10´ climate grids (Tracy and DeLoach, unpublished data), we estimate northern Uzbekistan as much more suitable habitat for D. carinulata than D. carinata. Therefore, we consider reports of D. elongata along the Amu Darya (river) in northern Uzbekistan by Yakhontov and Davletshina (1955, 1959), Sinadsky (1963, 1968) and Khamraev (2003) to probably refer primarily to D. carinulata rather than D. carinata (Map 5). Consequently, we consider the following four unconfirmed distribution records of D. elongata as uncertain records of D. carinulata (Map 5): UZBEKISTAN: Chertombayskoy Forest dachau [geocoordinates not locatable], Amu Darya Valley, heavy defoliation on Tamarix, VIII-1954; Nazarkhan [42.3333°N, 59.9833°E]; Nukus [42.8333°N, 59.4833°E]; Turtkul [41.5544°N, 61.0111°E] (Sinadsky 1963, 1968). Discussion. Taxonomy. Galeruca carinulata (Desbrochers 1870) was described from Sarepta, Russia and incorrectly synonymyzed under D. elongata var. carinata by Weise (1893). Berti and Rapilly (1973) removed D. carinulata from synonymy with both D. elongata and D. carinata based on their investigation of the endophallus. However, as discussed under D. carinata (above), only Warchalowski (2003) has accepted this taxonomic change. We dissected a single male topotype specimen (DEI 2003-07) from the type locality of D. carinulata in Sarepta, Russia, with endophalli matching that of Berti and Rapilly’s (1973) illustration of the endophallus of the holotype. This evidently old specimen (possibly from the late 1800’s) bore the following hand written label data: “Sarepta”; “G. carinulata Desbrochers”. A total of five males dissected from Astrakhan and Ikryanoye, Russia, ca. 365 km southeast of Sarepta (Map 5), also bore endophalli matching that of the description by Berti and Rapilly (1973), and three specimens of D. carinulata (of DEI) from Astrakhan also bore identification labels of G. carinulata Desbrochers. Three key characters of the endophallus of D. carinulata that distinguish it from other members of the D. elongata group can be seen in the illustrations of the endophallus of the holotype (Fig. 19b–c of Berti and Rapilly 1973) and our figures (Figs. 17, 22, 27, 32): (1) the acutely rounded distal margin of the palmate endophallic sclerite; (2) elongate endophallic sclerite with blade extending for less than 2/3 the length of the sclerite; and (3) lack of hooked apex of the elongate sclerite. Desbrocher’s (1870) holotype male for G. carinulata was 5.5 mm in length, below that usually found for D. carinata (mean male length is 6.29 mm, minimum male size is 5.12 mm, Table 2) but well within the size range of the 6 males we dissected from Sarepta, Astrakhan, and Ikryanoye (4.98–5.85 mm). We are confident that these specimens form a single species conspecific with D. carinulata. We find additional distinguishing characters in the endophallic sclerites and female genitalia (vaginal palpi and internal sternite VIII) of D. carinulata from the vicinity of the type locality in southern Russia and throughout its range to western China that distinguish it from D. elongata, D. carinata and other members of the D. elongata group (see Male- Genitalia and Female-Genitalia above; Figs. 17, 22, 27, 32, 37, 42; Map 5). The distinctive genitalic characters of D. carinulata are maintained in the same areas where D. carinata and D. elongata occur and near its abutting range boundary with D. meridionalis, and this is strong evidence for reproductive isolation

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 95 between these species (see Biogeography below; Map 1, Table 8). Although D. sublineata is allopatric with D. carinulata, their status as reproductively isolated is supported by the number of diagnostic genitalic characters separating D. sublineata from D. carinulata (6 characters) being greater than the number of characters separating the moderately sympatric D. carinata from D. carinulata (5 characters) (Table 5). Therefore, we firmly support Berti and Rapilly (1973) in restoring D. carinulata to species status, removing it from synonymy with both D. elongata and D. carinata. Habitus drawings of “D. elongata” adults from southern Russia (Fig. 30 of Ogloblin 1936) and Eastern Europe (Fig. 50 of Bieńkowski 2004) are probably D. carinulata. Drawings of “D. elongata” larval structures from southern Russia (Fig. 31 of Ogloblin 1936; Fig. 41.7 of Ogloblin and Medvedev 1971) probably also belong to D. carinulata. Chen (1961) described D. e. deserticola from the Taklamakan Desert region of Xinjiang Autonomous Region, China. The type locality of D. e. deserticola is Yuli (= Wei–li) (Map 5) and 8 additional localities are listed for paratypes. Yu et al. (1996) elevated D. e. deserticola to species status as D. deserticola without comment, and this change is mostly overlooked in recent biological literature (e.g., Li et al. 2000; DeLoach et al. 2003b; Lewis et al. 2003a, 2003b; Cossé et al. 2005; Milbrath and DeLoach 2006a), except by Meng et al. (2005) and Zhang and Baoping (2006). Lopatin et al. (2004) extended the range of D. e. deserticola to eastern Kazakhstan. We dissected two male and two female paratypes of D. e. deserticola collected from the type locality of Wei-lei, Xinjiang, China (Figs. 22, 27, 32 —Wei Lei Ta) and all belong to D. carinulata NEW SYNONYMY. We identified as D. carinulata all specimens dissected from Chen’s original given distribution of D. e. deserticola, China and Mongolia (Map 5). Chen’s description of D. e. deserticola gives ranges in body length of 5–6 mm for males and 5.5–7 mm for females, which is close to our observed size ranges of 4.6–6.1 mm for males and 5–7 mm for females (Table 2). Lewis et al. (2003b) gives mean lengths for males (5.6 ± 0.2 mm) and females (5.9 ± 0.2 mm) that are slightly larger than we observed in specimens collected over the entire range of D. carinulata (Table 2).The size range of 4.5–8 mm given for adult D. carinulata in regional keys of Mongolia (as D. e. ab. sublineata; Medvedev and Voronova 1977b, Medvedev 1982) and China (as D. e. sublineata; Gressitt and Kimoto 1963a) erroneously includes the larger size of other sibling species that can exceed 7 mm in length (Table 2), such as D. carinata, but which we have not found in these areas (Map1). An excellent habitus drawing of D. carinulata (as D. deserticola) from China is provided in Fig. 4-14 of Yu et al. (1996). Laboissière (1935) described D. koltzei ab. basicornis Laboissière from Khotan (=Hotan), Turkestan (now in Xinjiang Uygur Zizhiqu, China) from a single female. Wilcox (1971) listed D. k. ab. basicornis as a synonym of D. koltzei. However, Ogloblin (1936) had earlier synonymized D. koltzei under D. rybakowi and ranked it as the aberration D. rybakowi ab. koltzei. We dissected the genitalia of Laboissière’s (1935) female type specimen (from ZMAN) and identified it as D. carinulata. We also studied 9 syntypes of D. koltzei (from DEI; label data: Ili [Ili Kazakh Autonomous Prefecture, Xinjiang Uygur Zizhiqu, China] [18]’97; Weise; Coll. Koltze; Syntype) and 7 identified specimens of D. rybakowi (6 from USNM and 1 from DEI), and we concur that D. koltzei is conspecific with D. rybakowi which is very distinct from D. carinulata. Therefore, we conclude that D. k. ab. basicornis is conspecific with D. carinulata NEW SYNONYMY. The external characters provided by Berti and Rapilly (1973) to separate D. carinulata and D. carinata involve mainly the posterior angles of the pronotum and number of elytral carinae, and these characters are considered by Warchalowski (2003) as separating D. carinulata from D. elongata. However, we find these characters to be interspecifically variable and of no use in species diagnosis (see Discussion-Taxonomy under D. carinata). Chen (1961) noted that the ventral tarsal pubescence was medially absent, leaving a glabrous median line under the tarsi; in D. carinulata (as D. e. deserticola), but that the ventral tarsal pubescence is generally distributed in D. sublineata (as D. e. ab. sublineata) from North Africa. We examined dissected specimens of D. sublineata from North Africa with the same pattern of ventral tarsal pubescence as Chen described for D. e. deserticola, making this character too variable for use in species separation. We have dissected specimens of D. carinulata that were misidentified by taxonomists using external diagnostic characters as D. elongata from six countries and as D. e. sublineata or D. e. var. sublineata from Mongolia (see Material examined).

96 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Common Name. The vernacular name “striped tamarisk leaf beetle” has been applied to D. carinulata in Kazakhstan (as D. rybakowi; Mityaev 1958, Mityaev and Jashenko 2007) and China (as D. e. deserticola; Tian et al. 1988, Bao 1989, Sha and Yibulayin 1993). Because stripes can also be prominent in several other species of the D. elongata group (especially D. sublineata), we have dropped the term “striped” from the name and adopt the name “northern tamarisk beetle”, referring to its northernmost range of all tamarisk beetles (Fig. 51B, Map 1). Biology. Host Plants. Sha and Yibulayin (1993) listed the following hosts of D. carinulata (as D. e. deserticola) in order of preference based on numbers collected from the Turpan Eremophyte Botanical Garden in China: Tamarix laxa (Wildenow), T. elongata (Ledebour), T. kansuensis Zhang, T. gracilis Wildenow, T. androssowii Litvinov, T. arceuthoides, T. hispida, T. smyrnensis (as T. hohenackeri), and T. chinensis (Table 1). Mityaev and Jashenko (1998, 2007) reported T. leptostachya as a new host record from Kazakhstan among five other Tamarix spp. (see Table 1). The host range of D. carinulata (as D. e. deserticola) was reviewed by DeLoach et al. (2003b) who reported T. hispida var. karelinii (Bunge) Baum as a new host. DeLoach et al. (2003b) found the highest field populations on T. ramosissima among a total of seven Tamarix spp. on which it was collected in western China (including T. leptostachya, which was omitted from DeLoach et al. 2003b) (Table 1). Our collaborators R. Jashenko, I. Mityaev, and C. J. DeLoach collected D. carinulata from T. ramosissima at six locations and Tamarix sp. at three locations in Kazakhstan. Tamarix aralensis is a new host record for D. carinulata collected from Dry Sport Lake, Ashgabat, Turkmenistan by S. Myartseva (GSWRL collection). In addition to Tamarix spp., D. carinulata is feeds upon Myricaria alopecuroides Schrenk (Tamaricales: Tamaricaceae) in Mongolia (as D. elongata; Medvedev and Voronova 1979) and on one occasion was found to completely defoliate a few shrubs of Myricaria sp. by the Shelek River in southeastern Kazakhstan (as D. e. deserticola; Mityaev and Jashenko 1997, 2007). Sinadsky (1968) reported what is probably D. carinulata (as D. elongata, see above discussion) producing heavy defoliation on T. ramosissima compared to light damage on T. hispida along the Amu Darya in Karakalpakia [Qoraqalpog`iston Respublikasi], northern Uzbekistan. Tamarix kotschyi Bunge (as T. leptopetela Bunge) was recorded among the vegetation sampled at Abareq, Iran (Hoberlandt 1981) in the collection of at least 142 specimens of D. carinulata that we examined. Later detailed vegetation surveys in this area map T. kotschyi as the only Tamarix species in the vicinity of Abareq (Baum 1983), and we consider T. kotschyi a possible host for D. carinulata. Although T. aphylla is not a recorded host of D. carinulata, T. aphylla is common in the Registan-North Pakistan Sandy Desert in eastern Iran (Browicz 1991), an area from which we dissected D. carinulata from two locations (Map 6). Populations of D. carinulata established in North America are vigorously defoliating T. ramosissima in Nevada, Utah, Wyoming and Colorado and T. chinensis in Colorado (DeLoach et al. 2004). Large populations in western Nevada have fed upon cultivated T. parviflora, although this host is less preferred than T. ramosissima (Dudley et al. 2006, Dudley et al. in prep.) as Dalin et al. (in press) also observed in multiple-choice field cage studies in California. Tamarix parviflora is a new host record and novel host association in that it is not native where D. carinulata occurs in Asia. Attack on T. parviflora by D. carinulata is predictable from field cage studies in which T. parviflora did not significantly differ from most accessions of T. ramosissima/T. chinensis in terms of either suitability for larval survival or preference for adult oviposition (Milbrath and DeLoach 2006a). DeLoach et al. (2003b) found that D. carinulata (as D. e. deserticola) larvae from Turpan and Fukang, China and Shelek, Kazakhstan had highest survival on Tamarix followed by Myricaria and Frankenia. Adults oviposited less on bouquets of T. aphylla compared to T. ramosissima in the laboratory. Lewis et al. (2003a) found larval survival in the laboratory and adult oviposition in both laboratory and field cages on three North American Frankenia spp. (F. s a li na , F. johnstonii, and F. jamesii) generally were significantly lower than that on species of invasive North American Tamarix. But Milbrath and DeLoach (2006a) found that survival of larvae from Turpan was not different among the three North American Frankenia spp. and five invasive Tamarix spp. However, they confirmed previous results that adult oviposition was insignificant on the three Frankenia spp. in a combination of choice and no-choice field cage studies. Dudley and Kazmer (2005) planted alkali heath, Frankenia salina, at a site with large populations of D. carinulata defoliating tamarisk

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 97 near Lovelock, Nevada. In this open field test, D. carinulata did not lay eggs on the Frankenia and larvae crawling from defoliated tamarisk trees produced <4% leaf damage to Frankenia. In field cage tests, D. carinulata oviposited significantly less on T. aphylla compared to other invasive North American tamarisks, including T. ramosissima, T. chinensis, T. canariensis/T. gallica (Lewis et al. 2003a, Milbrath and DeLoach 2006a). Tamarix aphylla is at moderate risk of damage by D. carinulata in the field (DeLoach et al. 2003b), but the degree of damage that D. carinulata might cause to T. aphylla, especially in the absence of other Tamarix spp, is difficult to predict. Frankenia is at very low risk of damage from D. carinulata (Lewis et al. 2003a, Milbrath and DeLoach 2006a). Risk of damage to both T. aphylla and Frankenia by D. carinulata is probably much lower when these plants are not in the proximity of preferred Tamarix spp. (e.g., Blossey et al. 2001). Ecology. Diorhabda carinulata (as D. e. deserticola) is probably the most damaging specialized herbivore of tamarisk in Asia as a result of its abilities to: (1) defoliate large acreages of tamarisk, (2) reach high population densities, (3) produce several overlapping generations per season, (4) aggregate in large numbers, and (5) disperse widely from 2–6 km per day (Ding 2004). Sporadic outbreaks of D. carinulata (as D. e. deserticola or D. elongata) can defoliate 90% or more of the tamarisk over wide areas in Kazakhstan (DeLoach et al. 2003b), Uzbekistan (Sinadsky 1968), and China, where the beetles are controlled in order to protect tamarisk stands used in soil stabilization (Bao 1989, Tian et al. 1988, Sha and Yibulayin 1993, Chen et al. 2000, Ding 2004, Peng et al. 2005, Li and Wang 2006). In contrast, D. carinulata (as D. elongata) have been reported at population levels too low to damage tamarisk in the northern part of its range (ca. 45°N) in Mongolia (Medvedev and Voronova 1977b). Bao (1989) reported 66,666 ha of tamarisk colonized by D. carinulata (as D. e. deserticola), along the Donge He River (Ping Bao, Alashan Range Extension Station, Nei Mongol Zizhiqu, China, pers. comm.) of Erjina County, Inner Mongolia, China (Map 5). Of the 66,666 ha of colonized tamarisk, Bao reported 40,000 ha, or 60%, was defoliated by D. carinulata in this area (a region known for extensive stands of T. ramosissima; Kurschner 2004). Factors favoring periodic outbreaks of D. carinulata in Anxi County, Gansu Sheng, China, where up to 20,850 ha of tamarisk have been defoliated in a year, include (1) poor regulation from natural enemies, (2) mild winter temperatures increasing adult overwintering survival, (3) low precipitation during times of summer and fall pupation, reducing drowning of pupae on the ground, and (4) low water tables reducing mortality of ground dwelling pupae and overwintering adults (Ding 2004). Diorhabda carinulata extensively defoliated tamarisk stands in a 26,300 ha area over a 259 km stretch of the Humboldt River basin by late 2005 near Lovelock, Nevada, where it was introduced in 2001 (Carruthers et al. 2006, 2008). Large areas of tamarisk also have been defoliated near Lovell, Wyoming (D. Kazmer, pers. comm.), on the Colorado, Green, and Sevier rivers in Utah, and on the Dolores River in Colorado (D. Bean, pers. comm.). Smaller areas of tamarisk have been defoliated near Pueblo, Colorado (Map 7). Tamarisk defoliation by D. carinulata in Kazakhstan is primarily from feeding of first generation larvae in early summer (Mityaev and Jashenko 1999, 2007). Third instar larvae consume much more tamarisk than earlier stages in Ningxia Province, China (Tian et al. 1988). In Inner Mongolia, entire tamarisk trees are wilted from desiccation caused by larval feeding upon foliage and bark of tender branches, and tops of trees are discolored and wilted from aggregated feeding by adults (Chen et al. 2000). Densities per tamarisk bush reached over 10,000 larvae, and averaged 1,000 adults (maximum 4,000) in Inner Mongolia (Bao 1989). Near Turpan, China, average densities from 100 sampled tamarisk branches can reach 70 adults and 210 larvae per meter (Sha and Yibulayin 1993). Densities reach 270 adults and 534 larvae per meter of branch, and more than 7,000 larvae per individual tamarisk tree in Anxi County (Ding 2004). Densities of 173 to 247 larvae per meter of branch result in near defoliation of tamarisk near Sarytogay, Kazakhstan (Mityaev and Jashenko 1999, 2007). In northern Uzbekistan, larvae crawl from defoliated trees over ground to seek other tamarisk (Sinadsky 1968), a phenomenon that R. Carruthers and C.J. DeLoach also observed at Lovelock and at Schurz, Nevada, in 2004. Localized migrations of adult D. carinulata occur following emergence from overwintering, during mating of each generation, and prior to overwintering (Mityaev and Jashenko 1998, 2007).

98 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Diorhabda carinulata (as D. elongata) damages tamarisk over wide areas in all types of situations, including sand stabilization plantations and in the understory of turanga poplar ( Olivier), in northern Uzbekistan (Sinadsky 1968). Diorhabda carinulata more heavily damages the succulent tamarisk of riparian areas compared to that of drier upland areas in China (Bao 1989) and Kazakhstan (Mityaev and Jashenko 1998, 1999, 2007). Tamarisks growing in dense monoculture are attacked at a higher incidence compared to those in stands mixed with other shrubs such as Calligonum (Polygonaceae) and Haloxylon (Chenopodiaceae) near Turpan (Sha and Yibulayin 1993). Diorhabda carinulata defoliates tamarisk in the understory of plains cottonwood (Populus deltoides subsp. monilifera) at Pueblo, CO, and in extensive tamarisk monocultures at Lovelock, NV (C. Jack DeLoach, USDA/ARS, Temple, TX, pers. comm.). Tamarisk severely defoliated by D. carinulata completely resprouts in a short time in Kazakhstan (Mityaev and Jashenko 1998, 2007; see DeLoach et al. 2003b, Fig. 2). In tamarisk defoliated in the Chertombayskoy area of the Amu Darya, Uzbekistan, bud break is delayed by about 15 days the following spring and some of these bushes exhibit die back in the tops (Sinadsky 1968). Near Lovelock, Nevada regrowth also followed defoliation by D. carinulata, but it is usually accompanied by severe dieback and tree death is becoming more widespread in trees defoliated for several successive years (Carruthers et al. 2006, 2008). Tamarisk dieback and death at this site is probably related to observed significant reductions in nonstructural carbohydrates in tamarisk root crowns following one to four years of defoliation by D. carinulata (as D. e. deserticola) (Hudgeons et al. 2007b). Adult D. carinulata tend to aggregate in the field, and over 1000 adults were counted in a 2 x 2 m area in Inner Mongolia (Bao 1989). Near Chilik Kazakhstan, newly emerged first generation beetles aggregate on certain bushes just prior to mating (Mityaev and Jashenko 1999, 2007). In populations originating from Fukang, China, a male produced aggregation pheromone was identified consisting of two components: (2E,4Z)-2,4-heptadienal and (2E,4Z)-2,4-heptadien-1-ol (Cossé et al. 2005). This pheromone is being used to monitor populations near Lovelock, Nevada. A blend of green leaf volatiles is attractive for D. carinulata in the field, and the attraction is synergized when pheromone is added with the green leaf volatiles (Cossé et al. 2006). Additional observations on biology, including mating behavior, are noted by Tian et al. (1988), Bao (1989), Mityaev and Jashenko (1998, 1999, 2007), Zhang (2002), and Zhang and Baoping (2006) in central Asia. Phenology. Diorhabda carinulata has four generations from mid-April to mid-September at Turpan, China (Sha and Yibulayin 1993); three generations from April to October in northern Uzbekistan (Sinadsky 1968); three generations in Fukang (Li et al. 2000) and Ningxia province (Tian et al. 1988), China; two to three generations in Inner Mongolia Province, China (Bao 1989, Chen et al. 2000); and two generations in southern Kazakhstan (Mityaev and Jashenko 1998, 2007) and north of 38° in North America (DeLoach and Carruthers 2004b). Adults enter diapause from mid-August to September in China (Bao 1989, Tian et al. 1988, Sha and Yibulayin 1993). In southeastern Kazakhstan, adults begin overwintering in mid-September, but, if moisture is favorable and saltcedar has new growth, larvae still can be seen feeding on plants into September and adults are found into early October (Mityaev and Jashenko 1998, 2007). The typical hibernaculum of second generation adults overwintering in Kazakhstan is the boundary layer between the soil and detritus under saltcedar trees (Mityaev and Jashenko 1998, 2007). In Ningxia Province, China, adults overwinter under dead leaves or soil in areas facing the sun and protected from wind (Tian et al. 1988), At Lovelock, NV overwintering adults were found on the soil beneath the saltcedar litter during the fall and winter. Overwintered adults emerge in April in Ningxia Province (Tian et al. 1988) and in late April when temperatures rise over 13°C in Inner Mongolia (Bao 1989) and in May in southern Kazakhstan (Mityaev and Jashenko 1998, 2007). The Fukang, China D. carinulata climatype did not initially establish at several sites in North America south of 38°N latitude, probably due to asynchrony of the critical photoperiod inducing diapause with the onset of cooler temperatures in these areas (Bean et al. 2007a). In the south, these populations diapause prematurely in the summer, rather than the fall, leaving them inadequate food reserves for overwintering (Lewis et al. 2003b).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 99 Development and Reproduction. Diorhabda carinulata has three larval instars and a development time of 34 days from egg to adult at 24.1°C. Fecundity at 28.6°C on T. ramosissima averaged 194 eggs (range 78 to 550 eggs) with a population doubling time of 6.2 days. Fecundity on T. aphylla was lower than that found on T. ramosissima, T. canariensis and T. parviflora (Lewis et al. 2003b). Herrera et al. (2005) found 30–35°C is optimal among six constant temperatures from 15–40°C for highest survival and developmental rates for all life stages of D. carinulata. Both Zhang (2002) and Herrera et al. (2005) report temperature developmental thresholds and degree-days for development for various stages of D. carinulata. Milbrath et al. (2007) found that, at 28°C, D. carinulata (as D. elongata from Turpan and Fukang, China) had a development time of 18.5–20.4 days from egg to adult (with 74–78% survival), a fecundity of 272–283 eggs, and a population doubling time of 5.2–6.1 days. These values were all very similar to those found for D. elongata (Crete), D. carinata (Uzbekistan), and D. sublineata (Tunisia) (all as D. elongata). Natural Enemies. The parasitoid tachinid fly, Erynniopsis antennata, attacks larvae and emerges from adult D. carinulata from Fukang and Wujiaqu, Xinjiang Uygur Zizhiqu, China (Zhang 2002). At quarantine in Temple, Texas, we commonly encountered the parasitoid E. antennata emerging from the bloated abdomens of killed overwintered adult D. carinulata that were originally collected alive in April at Fukang, China. Diorhabda carinulata from Fukang, represents both a new host and locality record for E. antennata. Unidentified mymarid egg parasitoids attack as much as 24% or more of D. carinulata eggs in the field near Fukang and Wujiaqu, China (Zhang 2002, Zhang and Baoping 2006). An unidentified 1mm-long "fly", possibly an eulophid parasitoid hymenopteran, parasitized ca. 14.5–32.5% of the D. carinulata pupae in Inner Mongolia (Bao 1989). The ubiquitous fungal pathogen Beauveria bassiana (Balsamo) was found infecting adult D. carinulata originating from the following locations (all identified by T. Poprawski): Fukang, China (shipment GSWRL(CJD)-1996-27); near Urumqi, China (shipment GSWRL-1998-12); and near Shelek, Kazakhstan (shipment GSWRL-2000-9). At Turpan, China, the chief predators of D. carinulata are reduviids and mantids (Sha and Yibulayin 1993). Ants and coccinellids prey upon eggs and larvae of D. carinulata in Inner Mongolia (Bao 1989). In southeastern Kazakhstan, nymphs and adults of the pentatomid Arma custos Fabricius, feed on larvae of D. carinulata (Mityaev and Jashenko 2000, 2007). Seven species of birds have been reported to feed upon adult D. carinulata. In Inner Mongolia, tree sparrows, Passer montanus (Linnaeus), and common pheasants, Phasianus colchicus (Linnaeus), were observed eating adult beetles, and the crop of one common pheasant hunted in late winter contained over 200 overwintering adults along with a few wheat (Bao 1989). Chen et al. (2000) also noted the crested , Galerida cristata (Linnaeus), as a predator in Inner Mongolia. In Almaty, Kazakhstan, beetle adults were preyed upon by the great tit (Parus major Linnaeus), Eurasian blackbird (Turdus merula Linnaeus), common myna (Acridotheres tristis [Linnaeus]), and house sparrow (Passer domesticus [Linnaeus]) (Mityaev and Jashenko 2000, 2007). Biogeography. Comparative. Diorhabda carinulata differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) primarily continental, usually beyond 1,400 km from the ocean at elevations above 400 m; (2) commonly found in temperate cold desert and grassland biomes; and (3) latitudinal range of 29–49°N and most common at 40–44°N. Diorhabda carinata and D. carinulata are moderately sympatric and syntopic over portions of their western range (see above discussion under D. carinata-Biogeography) (Map 1, Table 8), and they are the most similar species in the D. elongata group in terms of biomes inhabited (Tables 9 and 10, Fig. 53). Diorhabda carinata differs from D. carinulata in being commonly collected in southern areas from 35–42°N and in preferring temperate grasslands over deserts. Diorhabda carinulata is marginally sympatric with D. elongata in southern Russia (Dagestan Republic) and allopatric with D. sublineata (Map 1; Table 8). Diorhabda elongata and D. sublineata both differ from D. carinulata in being primarily maritime and most common in the Mediterranean biome. Diorhabda carinulata is parapatric with D. meridionalis in southern Iran. Both D. carinulata and D. meridionalis have a similar preference for deserts, but D. meridionalis differs in being maritime and most common further south at 26–30°N (Maps 1 and 6; Figs. 51–52; Table 8).

100 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Descriptive. Diorhabda carinulata occurs over the central portion of the Palearctic realm (Maps 1, 5 and 6). It has been most commonly collected between 40–44°N in the Deserts and Xeric Shrublands and Temperate Grasslands and Shrublands biomes of eastern Central Asia. All reports of damage to tamarisk by D. carinulata are from this region of the Palearctic realm from elevations of ca. 30 m (at Turpan, China) to 970 m (Erjina County, Inner Mongolia, China). Diorhabda carinulata also occurs in Montane Grasslands and Shrublands biomes in western China around 44°N (from ca. 550–1850 m). From 35–40°N it is usually found in the Deserts and Xeric Shrublands biome from 0–1,400 m elevation. From 36–37°N, D. carinulata is occasionally found in the Temperate Conifer Forests biome in Iran (at ca. 1,400 m elevation). From 29–34°N in Iran, collections are from two biomes: Montane Grasslands and Shrublands from ca. 1,625–1,850 m elevation in the Kuh Rud and Eastern Iran Montane Woodlands ecoregion; and the Deserts and Xeric Shrublands biome from ca. 450–700 m elevation in the Registan-North Pakistan Sandy Desert (Map 6). The distribution of D. carinulata broadly coincides with that of the common host T. ramosissima from Mongolia and China to Iran and southern Russia, but its range appears to fall short of following the western distribution of T. ramosissima into Iraq and Turkey (Map 5). Primary ecoregions for D. carinulata in the northern portion of its range (40–46°N) in the Deserts and Xeric Shrublands include the Taklamakan Desert (30–1,600 m), Junggar Basin Semi-Desert (500–1,200 m), and Alashan Plateau Semi-Desert (900–1600 m) in China and Mongolia (Map 5). The primary northern ecoregion of D. carinulata in the Temperate Grasslands and Shrublands biome is the Tian Shan Foothill Arid Steppe (500–1,850 m) in southeastern Kazakhstan and northern Tajikistan. Potential in Tamarisk Biological Control. Summary. The northern tamarisk beetle is highly effective in biological control of T. ramosissima/T. chinensis in temperate cold deserts of the Great Basin Shrub Steppe, Colorado Plateau Shrublands and Wyoming Basin Shrub Steppe (Map 13). Diorhabda carinulata may be the most suitable tamarisk beetle for some warm temperate desert areas also, such as western portions of the Mojave Desert, southern portions of the Colorado Plateau Shrublands and Trans-Pecos Chihuahuan Desert (Map 13). Introduction of southern climatypes of D. carinulata from 29–34°N in the Registan North Pakistan Sandy Desert (Map 6) may speed its adaptation to corresponding latitudes in North America. Discussion. A northern D. carinulata climatype from ca. 44°N in China and Kazakhstan has been released and established in North America at several locations north of 37°N (Map 7). Diorhabda carinulata from Fukang, China, were introduced into cages in 1999, and released (under permit as D. elongata) into the field in 2001 at four sites in Nevada, Colorado, and Wyoming for biological control of T. ramosissima/T. chinensis. Concurrently, populations from Shelek, and Lavar (10 km west Shelek), and Buryndysu environs (25 km east Shelek), Kazakhstan, were released and established in Utah (DeLoach et al. 2004) (Maps 7, 9). In 2005, the USDA-APHIS (2005) began a large-scale introduction program of D. carinulata (as D. e. deserticola) from Fukang, China into 13 western states north of 38°N latitude. States involved in this program include Colorado, Idaho, Iowa, Kansas, Missouri, Montana, Nebraska, Nevada, North Dakota, Oregon, South Dakota, Washington and Wyoming. By 2005, it had defoliated over 20,000 ha of tamarisk in the Humboldt River basin near Lovelock, Nevada (Map 7). The northern tamarisk beetle has a moderate risk of damaging T. aphylla (DeLoach et al. 2003b) and a low risk of damaging Frankenia (Lewis et al. 2003a, Milbrath and DeLoach 2006a), and both these risks are probably much reduced at less proximity to preferred Tamarix spp. (e.g., Blossey et al. 2001). The introduced northern D. carinulata climatype probably has the greatest potential to defoliate tamarisk in North America in the temperate cold Deserts and Xeric Shrublands and the Temperate Grasslands and Shrublands biomes from 40–44°N (Map 13, see Biogeography). Elevations above 1,000 m might extend this defoliation further south in some ecoregions. Diorhabda carinulata has already defoliated large acreages of tamarisk at sites in three North American temperate cold desert ecoregions: the Great Basin Shrub Steppe (Lovelock, NV and Delta, UT), the Colorado Plateau Shrublands (Potash, UT), and the Wyoming Basin Shrub Steppe (Lovell, WY) (Map 13). The southern portion of the Snake/Columbia Shrub Steppe is probably also a highly suitable cold desert region (Map 13). The most suitable ecoregions of the Temperate Grasslands and Shrublands biome at 40–44°N probably include the southern portion of the Northwestern Mixed Grasslands

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 101 and the northern portions of the Western Short Grasslands (Map 13). Diorhabda carinulata has also established well at 38°N at Pueblo, Colorado in the Western Short Grasslands, but D. carinata may be better suited to the temperate grassland biome at this latitude (Map 13). The potential spread of the introduced parasitoid tachinid fly Erynniopsis antennata from California to Nevada should be monitored as it might severely reduce overwintering survival of northern tamarisk beetles. The northern climatype of D. carinulata from Fukang, China was initially ill-adapted to areas south of 38°N (DeLoach et al. 2004) because of premature entry into diapause during short summer daylengths (Lewis et al. 2003b; Bean et al. 2007a, 2007b). Overwintering of the northern Fukang climatype failed at field cages in Big Pine, California, Artesia, New Mexico, and Seymour, Temple, and Lake Thomas, Texas. The lower elevation D. carinulata northern climatype from Turpan, China failed to establish in cages at Lake Thomas, Texas (personal observation), and is establishing poorly at John Martin Reservoir, Colorado (Debra Eberts, USDI/Bureau of Reclamation, Denver, CO, pers. comm.) (Map 7). However, in 2008, populations originating from Delta, Utah, had established south to about 37°N on the Virgin River at Littlefield, Arizona (L.D. Walker, pers. comm.) and in Ute Canyon, Colorado (Dan Bean, pers. comm.). Natural expansion of the population was rapid in 2008, covering over 75 km south of Gateway along the Dolores River (D. Bean, pers. comm.). In 2006, a population of the northern Fukang climatype from Nevada began to increase and defoliated about 0.8 ha of tamarisk in the open field at Artesia, New Mexico (at ca. 33°N), but populations disappeared by 2008 (D. Thompson, pers. comm.). Apparently, the northern Fukang climatype may be gradually adapting further south. Putative climatypes from the extreme southern range limits of D. carinulata occur from 29–34°N in widely varied habitats of southeastern Iran, such as the Registan-North Pakistan Sandy Desert (at ca. 450–700 m elev.) and Kuhrud Mountains (at ca. 1,625–1,850 m elev.) (Map 6). In North America, these southern climatypes may more quickly adapt to areas in the southern half of the Colorado Plateau Shrublands, western portions of the Mojave Desert, Trans-Pecos Chihuahuan Desert, and southwestern portions of the Western Short Grasslands ecoregions. According to our HSI models, D. carinulata will probably not establish further south in subtropical deserts such as the Sonoran, Tamaulipan Mezquital and southern Chihuahuan (Map 13). Initial attempts at establishing the northern Fukang D. carinulata climatype were unsuccessful north of 46°N at Fort Peck, Montana (D. Kazmer, pers. comm.) (Map 7). Putative climatypes of D. carinulata from the extreme northern limits of its native range occur from 46–48°N at low elevations (ca. 0 m) in the Volga Valley of southern Russia and higher elevations (at ca. 1,200 m) of the Junggar Basin Semi-Desert in western Mongolia (Map 5). These extreme northern climatypes of D. carinulata may be better preadapted to Montana.

Diorhabda meridionalis Berti & Rapilly, 1973 NEW STATUS southern tamarisk beetle (Figs. 9, 18, 23, 28, 33, 38, 43)

Diorhabda carinulata meridionalis Berti & Rapilly, 1973:881 (Type locality: Minab, Iran); Warchalowski, 2003:328 (catalog of Mediterranean Region, Iran). Diorhabda elongata: Riley et al., 2003:69,189 (part, catalog of North America [not yet introduced]).

Male. Genitalia. Diorhabda meridionalis can be distinguished from all other members of the D. elongata group by two unique characteristics of the elongate endophallic sclerite (EES): (1) the presence of a strongly hooked apex (Figs. 18, 28), and (2) the length of the blade of the EES being greater than or equal to 0.67 times the length of the EES (Table 3; Figs. 18, 23, 48). In all other members of the D. elongata group, the blade is less than or equal to 0.66 times the length of the EES (Figs. 14–17, 19–22) and the apex of the EES is not hooked (Figs. 24–27). The palmate endophallic sclerite (PES) of D. meridionalis is also unique among the D. elongata group in that its distal margin is narrowly rounded and generally smooth, with only one or two small subdistal spines, and it has a length to width ratio of 0.35–0.63 (Figs. 18, 33; Table 3). In contrast, the distal margin of the PES is truncate or more broadly rounded with larger distal or subdistal spines in D. elongata, D.

102 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS carinata, and D. sublineata (Figs. 14–16, 29–31), and the PES is acutely rounded with a length to width ratio of 0.61–1.02 in D. carinulata (Figs. 17, 32). Measurements. See Tables 2 and 3. Female. Genitalia. Female D. meridionalis can be distinguished from all other members of the D. elongata group by the following combination of characters in the vaginal palpi (VP) and internal sternite VIII (IS VIII): (1) the vaginal palpi are broadly rounded and as long as wide or longer with a width to length ratio (LP/WP) of 1.0–1.31 (Fig. 38; Table 4), and (2) the width of the stalk (WST) of IS VIII is 0.22–0.33 mm and this width is 0.33–0.48 times the width of the apical lobe (WAL) (Figs. 38, 43; Table 4). In contrast, the vaginal palpi are wider than long, with a length to width ratio of 0.46–0.94 in D. elongata, D. carinata and D. sublineata (Figs. 34–36). In addition, the vaginal palpi are triangulate in D. carinata and D. sublineata (Figs. 35–36). In D. carinulata, the width of the stalk of IS VIII is 0.36–0.57 mm and this width is from 0.49–0.77 times the width of the apical lobe (Fig. 37, 42). In addition, the width of the widest lobe of the stalk of IS VIII is from 0.04–0.09 mm in D. meridionalis (Fig. 38, 43); whereas, in D. carinulata the width of the widest lobe of the stalk is usually greater than 0.10 mm (Figs. 37, 42; Table 4). Measurements. See Tables 2 and 4. Coloration. Coloration of dead D. meridionalis (Fig. 9) is similar to that discussed above for D. sublineata and D. carinulata (Figs. 5, 7). We have not had the opportunity to examine live material of D. meridionalis. Type material. Berti and Rapilly (1973) deposited the holotype male and allotype female of D. meridionalis, both from Minab, Iran, and four paratypes (3♂♂, 1♀) from Minab, Borazjan and Shush (= Susa), Iran at MNHN. As discussed above (see D. elongata - Type Material), we were unable to obtain type materials from MNHN. We studied the original description by Berti and Rapilly (1973), including their illustrations of the endophallus of the male holotype. We also examined a male specimen from 30 km NNE of Borazjan, a paratype locality, with an identification label as “D. carinulata meridionalis Berti & Rapilly det. Berti X-1996” (MBPF 2003-03) and additional material near type and paratype localities. Material examined. 22♂♂ dissected (diss.), 19♀♀ diss., 13♂♂, 18♀♀, 147 unsexed specimens. IRAN: 2♂♂ diss., 3♀♀ diss., 11♂♂, 9♀♀, Bahu–Kalat [25.73330°N, 61.53330°E; 68 km south Rask, Sarbaz River; T. dioica and T. aphylla present (Hoberlandt 1981)], 3–4-IV-1973, Exp. Nat. Mus. Praha Loc. No. 147, NMPC [2004-48, 2005-16, 17, 18, 19]; 3♂♂ diss., Bilai [26.48333°N, 57.15000°E; coastal savanna; Tamarix sp. not recorded (Hoberlandt 1981)], 23–24-V-1973, Exp. Nat. Mus. Praha Loc. No. 209, Diorhabda elongata det. I.K. Lopatin 1978 [1♂ diss., USNM], NMPC [2004-57, 2005-22], USNM [2003-11]; 1♂ diss., 2♀♀ diss., 1♀, Borazjan, 20 km northwest [29.38333°N, 51.03333°E; Shabankareh, clay semi-desert; Tamarix sp. not recorded (Hoberlandt 1983)], 18-IV-1977, Exp. Nat. Mus. Praha Loc. No. 297, NMPC [2004-05, 42, 43]; 3♂♂ diss., 1♂, 3♀♀, Borazjan, 30 km north northeast [29.53333°N, 51.40000°E; 10 km N Dalaki, Hableh Rud River; Tamarix sp. present (Hoberlandt 1983)], 18–19-IV-1977, Exp. Nat. Mus. Praha Loc. No. 298, D. carinulata meridionalis Berti & Rapilly det. Berti X-1996 [1♂ diss., MBPF], NMPC [2004-12, 2005-20], MBPF [2003-03]; 2♀♀ diss., Jeiugir (33° 27' N, 49° 01' E) [sic; Jolow Gir; 32.96667°N, 47,81611°E; coordinates corrected from relation to Sarab–e Jahangir], 20 km southeast Sarab–e Jahanagir, Lorestán, 8–10- X-1998, Chvojka, D. elongata det. A. Warchalowski, NMPC [2005-28, 2006-01]; 1♂ diss., 2♀♀ diss., 1♀, Nikshahr, 13 km south southeast [26.13333°N, 60.18333°E; Rudkhaneh Nikshahr river; T. dioica present (Hoberlandt 1981)], 8–9-IV-1973, Exp. Nat. Mus. Praha Loc. No. 152, USNM [2003-05, 2004-05, 06], NMPC; 1♂ diss., 1♂, 3♀♀, Omidiyeh, 34 km southeast [30.55000°N, 49.91667°E; Rud-e Zohreh river; Tamarix sp. present (Hoberlandt 1983)], 16–17-IV-1977, Exp. Nat. Mus. Praha Loc. No. 292, NMPC [2004- 44]; 8♂♂ diss., 5♀♀ diss., 1♂, 2♀♀, 142 specimens [NMPC], Rask [26.23528°N, 61.40111°E; 3 km N Rask; T. dioica present (Hoberlandt 1981)], Sarbaz River valley, 3–4-IV-1973, Exp. Nat. Mus. Praha Loc. No. 146, D. carinulata meridionalis det. Lopatin 1984 [1♂, 1♀, NMPC], D. elongata det. I.K. Lopatin [2♀♀, USNM], USNM [2003-04, 12, 13, 2004-03; 2008-02, 03, 04, 05, 06], NMPC [2004-58, 2005-21, 2006-8–9]; 1♂ diss., 2♀♀ diss., 1♀, Sabzevaran [Sabzvaran], 33 km south [28.40000°N, 57.83333°E; ruderal plants near plantation; Tamarix sp. present (Hoberlandt 1983)], 17-V-1977, Exp. Nat. Mus. Praha Loc. No. 335, NMPC

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 103 [2004-27, 2005-23, 24]; 1♀ diss., Sekand [26.71667°N, 61.53330°E; oasis of saline marshes; Tamarix sp. not recorded, Frankenia pulverulenta present (Hoberlandt 1981)], 27 km east northeast Sarbaz, 31-III to 1-IV- 1973, Exp. Nat. Mus. Praha Loc. No. 144, USNM [2004-02]; 1♀ diss., Shushtar [32.05°N, 48.85°E; river Karun; Tamarix sp. present (Hoberlandt 1983)], 13-IV-1977, Exped. Nat. Mus. Praha Loc. No. 287, NMPC [2005-35]; PAKISTAN: 1♂ diss., 1♀ diss., Turbat [26.02780°N, 63.05060°E], west Balochistan, 8–19-IV- 1993, S. Becvar, EGRC [2005-01, 04]; SYRIA: 1♂ diss., Alep[po] [Halab; 36.20278°N, 37.15861°E], Syrie, NHMB [2003-15]. Distribution. General. Diorhabda meridionalis is primarily known from extreme southern Iran and along the southern portion of Iran’s border with Iraq. Its native range is from Syria to western and southern Iran and southern Pakistan (Map 5). Additional surveys should be made for D. meridionalis in eastern Iraq and along the western coast of the Persian Gulf in Kuwait, Saudi Arabia, , Oman, and east to northern India (following the distribution of a suspected host T. dioica Roxburgh ex Roth, see below; Map5). Confirmed Records. We confirm the presence of D. meridionalis in Iran (Berti and Rapilly 1973) (see above Material examined) (Maps 5–6). Although D. meridionalis, D. carinulata and D. carinata occur on tamarisk throughout Iran (Map 6), Barkhodari et al. (1981) recorded no Diorhabda in their detailed survey of the Tamarix entomofauna in Iran. This is evidence of a possibly sporadic nature of tamarisk beetle populations in Iran. New Records. Pakistan and Syria are new distribution records for D. meridionalis (Map 5). Unconfirmed Records. We find no country distribution records that remain unconfirmed. We have not examined specimens from the following locations of D. meridionalis in Iran reported by Berti and Rapilly (1973) (Map 5): IRAN: 1♂, 1♀, Borazjan [29.26222°N, 51.20528°E], 24-V-1969, R. Naviaux and M. Rapilly, on Tamarix sp., PARATYPE, MNHN; 2♂, 1♀, Minab [27.14667°N, 57.07361°E], 25-IV-1971, Naviaux and M. Rapilly, on Tamarix sp., HOLOTYPE, MNHN; 1♂, Susa [Shush; 32.19417°N, 48.24361°E], 1899, Escalera, PARATYPE, MNHN. Discussion. Taxonomy. Diorhabda carinulata meridionalis was described from Minab, Iran with paratype localities in Shush and Borazjan, all along the southwestern border of Iran (Berti and Rapilly 1973). We examined one male specimen from 30 km NNE of Borazjan, a paratype locality, with an identification label as “D. carinulata meridionalis Berti & Rapilly det. Berti X-1996” (MBPF 2003-03) with endophalli matching that of the description by Berti and Rapilly (1973). Other males with endophalli matching that of D. c. meridionalis were examined from Bilai, 74 km NNW from the type locality of Minab, from 20 km NNW of Borazjan, and from six other locations along the southwestern Iranian border (Map 5). Three key characters of the endophallus of D. c. meridionalis that distinguish it from other members of the D. elongata group can be seen in the illustrations of the endophallus of the holotype (Fig. 19d–e of Berti and Rapilly 1973) and our figures (Figs. 18, 23, 28, 33): (1) narrowly rounded distal margin of the palmate endophallic sclerite; (2) elongate endophallic sclerite (EES) with blade extending for more than 2/3 the length of the sclerite; and (3) the hooked apex of the EES. We are certain that specimens we studied with endophalli matching that of D. c. meridionalis are conspecific. We find additional characters in the endophallic sclerites (Figs. 18, 23, 28, 33) and female genitalia (vaginal palpi and internal sternite VIII; Figs. 38, 43) of D. c. meridionalis throughout its range (Map 5) that distinguish it from other members of the D. elongata group (see Male-Genitalia and Female-Genitalia above; Figs. 14–17, 19–22, 24–27, 29–32, 34–37, 39–42). If D. c. meridionalis were a subspecies of D. carinulata, we would expect intermediate forms to occur in areas where their distributions approach one another in southern Iran. Diagnostic characters of the endophalli and female vaginal palpi of D. c. meridionalis are distinct from that of D. carinulata, even in the populations at Sabzvaran, Iran, that approach within 102 km of D. carinulata at Abaraq, Iran (see Biogeography below; Maps 1 and 5, Table 8). The lack of intermediate forms between the two taxa is strong evidence of their reproductive isolation. The distinctive genitalic characters of D. c. meridionalis are also maintained in the same areas where D. carinata occurs and near the abutting range boundary of D. c. meridionalis with D. elongata, and this is strong evidence for reproductive isolation between these species. Although D. c. meridionalis is allopatric with D.

104 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS sublineata, their status as reproductively isolated is supported by the number of diagnostic genitalic characters separating D. c. meridionalis from D. sublineata (8 characters) being greater than the number of characters separating the moderately sympatric D. carinata from D. carinulata (5 characters) (Table 5). Therefore, we elevate D. c. meridionalis to species status as D. meridionalis Berti and Rapilly NEW STATUS. We were unable to fully evert the tip of the endophallus containing the gonopore in D. meridionalis without damaging the specimens (Fig. 18), and a more fully everted pointed tip can be seen in the illustration of the holotype endophallus in Fig. 19d–e of Berti and Rapilly (1973). We have examined specimens of D. meridionalis that were misidentified by taxonomists as D. elongata from Iran (see Material Examined).

FIGURES 44–47. Palmate, connecting (dorsal views), and elongate (dorsal and lateral views) endophallic sclerites with abnormal sclerites in laboratory pure lines of Diorhabda elongata and D. sublineata and laboratory produced D. sublineata × D. elongata hybrids from New Mexico State University (NMSU) . 44—♂ D. elongata pure line, 45—♂ D. sublineata pure line, 46—♀ D. sublineata × ♂ D. elongata hybrids (F1 and F2), 47—♀ D. elongata × ♂ D. sublineata hybrids (F1 and F2). AS—abnormal sclerite, CES—connecting endophallic sclerite, DS—distal spines, EES—elongate endophallic sclerite, LA—lateral appendage, LB—length of blade, LN—lateral notch, PES—palmate endophallic sclerite, SDS—subdistal spines, SL—length of spined area of blade. Scale bar 1.0 mm.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 105 FIGURE 48. Scatter plots of length of blade of elongate endophallic sclerite (A) and length of spined area of elongate endophallic sclerite (B) versus the length of the elongate endophallic sclerite for field collected material of Diorhabda elongata species group and laboratory produced F1 and F2 D. elongata/D. sublineata hybrids (see Figs. 45–47 for illustrations of endophallic sclerites for individual hybrids identified by alphanumeric codes in plots; see Table 3 for sample sizes and statistics for measurements).

106 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURES 49–50. Genitalic phenograms for the Diorhabda elongata species group. 49–dissimilarity phenogram based on average taxonomic dissimilarity matrix (Table 5), 50—similarity phenogram based on and Pearson product-moment correlation similarity matrix (Table 6). Phenograms produced with NTSYSpc Tree plot module (Rohlf 2006) from clusters formed with unweighted arithmetic average clustering (UPGMA) (NTSYSpc SAHN module).

Common Name. The vernacular name “southern tamarisk beetle” refers to its Latin name “meridionalis” and its southern range in relation to the closely related and parapatric D. carinulata (Fig. 51B). Biology. Host Plants. Berti and Rapilly (1973) report Tamarix sp. as a host plant for D. meridionalis (as D. c. meridionalis) in Iran (Table 1). Records of Tamarix identified to species and other plants present were made in conjunction with collections of D. meridionalis in southeastern Iran by the second Czechoslovak- Iranian Entomological Expedition in 1973 (Hoberlandt 1981; see above Material Examined). We examined D. meridionalis from one of two expedition collection sites with both T. dioica and T. aphylla (Bahu–kalat), two of five sites with T. dioica and no T. aphylla (Rask and 13 km SSE Nikshahr), and none of eight sites with T. aphylla and no T. dioica. Based on these records, T. dioica, a close relative of T. aphylla, is a probable host plant in southeastern Iran. We found no reports of T. dioica in western Iran and Syria (Schiman-Czeika 1964,

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 107 FIGURE 51. Schematic box plots for elevations (A) and latitudes (B) from native presence-only field collection data for the Diorhabda elongata species group at 5 minute grid resolution. Box plots for each species depict the mean, median, first quartile (Q1), third quartile (Q3), interquartile range (IQR, Q1–Q3), low whisker (LW; lowest point at or above 1.5*IQR lower than Q1), high whisker (HW, highest point at or below 1.5*IQR higher than Q3), mild outliers (MO, points between the low or high whisker and 3*IQR from Q1 and Q3, respectively), and extreme outliers (EO, points below or above 3*IQR from Q1 and Q3, respectively) (Proc Boxplot; SAS Institute 2005). Sample sizes of field localities (N) and a table summary statistics and plotted values are inset in each chart.

108 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURE 52. Schematic box plots for distances to the ocean (A) (Proc Boxplot [Boxstyle=Schematic]; SAS Institute 2005) and a bar chart of frequency percentages for distribution of each Diorhabda species across biomes (B) (Proc Freq; SAS Institute 2005) from native presence-only field collection data for the Diorhabda elongata species group at 5 minute grid resolution. Percentage of a species in a biome is number of collections for that species in the biome divided by the number of collections of that species across all biomes. See Figure 51 for explanation of box plots. Sample sizes of field localities (N) and a table summary statistics and plotted values are inset in each chart.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 109 FIGURES 53–54. Biomic dissimilarity dendrograms based on biomic Bray-Curtis dissimilarity matrices. 53—Diorhabda elongata species group (from Table 10), 54—both the D. elongata group and Tamarix species invasive in North America (from Table 12). Dendrograms produced with NTSYSpc Tree plot module (Rohlf 2006) from clusters formed with unweighted arithmetic average clustering (UPGMA) (NTSYSpc SAHN module). Line connecting D. elongata and T. gallica at left signifies that the positions of these taxa are interchangeable in an alternate dendrogram of equal rcoph value.

Baum 1978, Zieliński 1994), and other Tamarix spp. probably serve as hosts of D. meridionalis in these areas (Maps 5 and 6). Tamarix aphylla is common in southern Iran and southern Pakistan (Browicz 1991) where it might serve as a host for D. meridionalis. At the time of collection of D. meridionalis in an oasis of small saline marshes at Sekand, Iran, Hoberlandt (1981) recorded as present the annual herb Frankenia pulverulenta Linnaeus (of the order Tamaricales with Tamarix) but not Tamarix. Frankenia pulverulenta is indigenous to the southern Palearctic

110 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS region and southern Africa (Jäger 1992), and it sometimes grows in association with Tamarix in north Africa (Kassas and 1954), southern Europe (Cano et al. 2004), and probably also in Iran. It is unlikely that this herbaceous Frankenia would serve as a host of D. meridionalis, but in future surveys for D. meridionalis in the region, F. pulverulenta should be surveyed in order to rule it out as a host. Diorhabda meridionalis and D. carinata have been collected together in series from five locations in western Iran and Syria, and they probably share some of the same host Tamarix spp. in a manner similar to D. carinulata and D. carinata. Ecology and Phenology. Collection dates in Iran and Pakistan for D. meridionalis from examined material and collections records of Berti and Rapilly (1973) are from 31 March through 10 October. Natural Enemies. We found no reports of natural enemies of D. meridionalis.

FIGURE 55. Three dimensional biomic principal coordinate analysis (PCoA) scatter plot for the Diorhabda elongata species group and Tamarix spp. invasive in North America for the first three eigenvectors (cumulative axis loading of 80.92%; Table 13) computed from a biomic Bray-Curtis dissimilarity matrix (Table 12). Biomes in which ranks of species are statistically significantly positively or negatively correlated with ranks of species in each PCoA axis are indicated in parentheses (see Table 13). Plotted with Mod3D module of NTSYSpc (Rohlf 2006).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 111 FIGURE 56. Linear models for latitudinal (A) and elevational (B) habitat suitability indices for the Diorhabda elongata species group. Model parameters calculated from descriptive statistics (Fig. 51), including interquartile range (IQR), minimum (MIN) value minus 1% or 10% of range (R), and maximum value (MAX) plus 1% or 10% of range (labeled for D. elongata).

112 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS FIGURE 57. Linear and categorical models for continentality (A) and biomic (B) habitat suitability indices for the Diorhabda elongata species group. Model parameters calculated from descriptive statistics (Fig. 52), including interquartile range (IQR), minimum (MIN) value minus 20% of range (R), and maximum value (MAX) plus 20% of range (labeled in A for D. carinulata).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 113 FIGURE 58. Categorical model for biomic relative suitability indices for the Diorhabda elongata species group. Model parameters calculated from descriptive statistics (Fig. 52B).

Biogeography. Comparative. Diorhabda meridionalis differs from other tamarisk beetles by the following combination of biogeographic characteristics: (1) primarily maritime and found within 300 km of the ocean at elevations usually under 600 m (ranges to 1,100 m); (2) usually found in subtropical desert and temperate broadleaf and mixed forest biomes; and (3) latitudinal range of 26–36°N and most common at 26–31°N. Diorhabda meridionalis is partially sympatric with D. carinata, their ranges overlapping in western Iran and Syria where they appear to be syntopic (see above discussion under D. carinata—Biogeography) (Maps 1 and 6; Table 8). Diorhabda meridionalis is parapatric with D. carinulata in southern Iran (Map 6). Diorhabda meridionalis is most similar to the species pair D. carinata/D. carinulata in terms of biomes inhabited (Tables 9 and 10, Fig. 53). Both D. carinata and D. carinulata differ from D. meridionalis in being primarily continental in distribution and more common north of 31°N. Diorhabda meridionalis is allopatric with D. elongata and D. sublineata (Map 1, Table 8). Diorhabda elongata and D. sublineata differ from D. meridionalis in their greater preference of the Mediterranean biome and their commonness north of 31°N. Further collections might reveal that D. meridionalis and D. sublineata are parapatric or marginally sympatric in the eastern Saudi Arabian peninsula. Descriptive. Diorhabda meridionalis is a south central Palearctic species that may also occur at the northern tip of the Afrotropical realm in the United Arab Emirates (Map 1). It is most commonly collected from 26–31°N in the Deserts and Xeric Shrublands biome of southern Iran in the South Iran Nubo-Sindian Desert and Semi-Desert ecoregion (20–390 m) (Map 6). According to the biogeographic classification of Morrone (2002), more than half of this ecoregion falls within the northwestern tip of the Oriental region (= Indo-Malayan realm of Olson and Dinerstein [2002]) which is part of the Holotropical kingdom. But we follow Olson and Dinerstein (2002), whose placement of the western border of the Indo-Malayan and Palearctic realms near the border of Pakistan and India (Map 1) is supported by borders of the corresponding

114 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 1. Native distribution of the Diorhabda elongata species group. Some symbols for Diorhabda spp. overlap and some locations are approximate (see Maps 2–5).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 115 MAP 2. Native distribution of Diorhabda elongata with occurrences of selected Tamarix spp.

116 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 3. Native distribution of Diorhabda carinata with occurrences of selected Tamarix spp.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 117 MAP 4. Native distribution of Diorhabda sublineata with occurrences of selected Tamarix spp.

118 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 5. Native distribution of Diorhabda carinulata and D. meridionalis with occurrences of selected Tamarix spp.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 119 MAP 6. Native distribution of the Diorhabda elongata species group in part of southwest Asia with occurrences of selected Tamarix spp. Some symbols for Diorhabda spp. overlap and some locations are approximate (see Maps 2–5).

120 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 7. Distribution of the Diorhabda elongata species group where introduced in North America with occurrences of selected Tamarix spp. Some symbols for Diorhabda and Tamarix spp. overlap.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 121 MAP 8. Hand-fitted habitat suitability index (HSI) models for the Diorhabda elongata species group over their native distribution: A—Diorhabda elongata, B—D. carinata,C—D. sublineata.

122 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 9. Hand-fitted habitat suitability index (HSI) models for the Diorhabda elongata species group over their native distribution: A—Diorhabda carinulata, B—D. meridionalis. faunistic kingdoms of Bobrov (1997). The distribution of D. meridionalis in the South Iran Nubo-Sindian Desert and Semi-Desert ecoregion corresponds to the western distribution of the suspected host T. dioica,and further surveys might reveal more overlap between D. meridionalis further east and north along the distribution of T. dioica in Pakistan and India (Map 5). Habitats of collection sites for D. meridionalis in southern Iran are primarily river banks, but also include saline marshes, coastal savannas, clay semi-deserts, and weedy fields (Hoberlandt 1981, 1983). Diorhabda meridionalis is also found in the Mediterranean Forests, Woodlands and Scrub biome in Syria (36°N, ca. 380 m), the Temperate Broadleaf and Mixed Forests biome of western Iran (29°N; ca. 60–500 m) and the Montane Grasslands and Shrublands of southern Iran (27°N) to ca. 1,102 m elevation at Sekand (Table 9). Most collection sites are within 100 km of the sea, the maximum distance being 309 km at Jolow Gir, Iran (Table 7; Map 5).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 123 MAP 10. Hand-fitted habitat suitability index (HSI) models for the Diorhabda elongata species group over western North America: A—Diorhabda elongata, B—D. carinata,C—D. sublineata (See Maps 7 or 13 for legend of symbols for sites of Diorhabda establishment, release, or planned release).

124 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 11. Hand-fitted habitat suitability index (HSI) models for the Diorhabda elongata species group over western North America: A—Diorhabda carinulata, B–D. meridionalis (See Maps 7 or 13 for legend of symbols for sites of Diorhabda establishment, release, or planned release).

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 125 MAP 12. Composite map of habitat suitability index (HSI) models scoring among the top 15% among the Diorhabda elongata species group over their native distribution. The composite map depicts the estimated most suitable Diorhabda species or group of species for a given area, not the total potential range for any given species. Some symbols for Diorhabda spp. overlap and some locations are approximate (see Maps 2–5).

126 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS MAP 13. Composite map of habitat suitability index (HSI) models scoring among the top 15% among the Diorhabda elongata species group over North America. The composite maps depict the estimated most suitable Diorhabda species or group of species for a given area, not the total potential range for any given species. Some symbols for Diorhabda spp. overlap.

Potential in Tamarisk Biological Control. Summary. The southern tamarisk beetle may potentially be the best suited Diorhabda species for biological control of tamarisk in portions of maritime subtropical deserts such as the Sonoran Desert and Tamaulipan Mezquital (Map 13). Overseas collections of D. meridionalis may

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 127 be obtained from the South Iran Nubo-Sindian Desert and Semi-Desert in southern Iran and southwest Pakistan (Map 6). The preference of D. meridionalis for T. ramosissima/T. chinensis versus T. aphylla will probably be a critical issue for host range testing. In addition, the potential effects of D. meridionalis on Frankenia spp. are not known. Discussion. We find no reports of damage to tamarisk by the southern tamarisk beetle. This species can be abundant in areas such as Rask, Iran from which we examined a large series of at least 159 beetles collected in 1973 by the Czechoslovak-Iranian entomological expedition in an area where Hoberlandt (1981) recorded the presence of T. dioica. Diorhabda meridionalis probably attacks several Tamarix spp., as do other species in the D. elongata group, and it may also attack Tamarix introduced into North America. Host range studies are needed to verify that D. meridionalis is restricted in host range to Tamarix spp. and determine its preference for various Tamarix spp. introduced into North America. The Palearctic annual herb Frankenia pulverulenta, a close relative of Tamarix found at a collection site with D. meridionalis in Iran (see above), should be included in host range studies. Frankenia pulverulenta is naturalized in the U.S. states of Utah and Oregon (Welsh et al. 1987, USDA Natural Resources Conservation Service 2008). Maritime subtropical Deserts and Xeric Shrublands from 26–31°N may be most suitable for D. meridionalis in North America (Maps 5–6). Suitable North American ecoregions include the southern Sonoran Desert, the eastern Tamaulipan Mezquital, the Baja California Desert, and the Gulf of California Xeric Scrub (Map 13).

Implications Regarding Biological Control of Tamarisk

Our study illustrates that basic research into the taxonomy and biogeography of potential biological control agents can be essential to timely and effective implementation of weed biological control programs. Below is an outline of our major conclusions as they relate to biological control of tamarisk.

1) Five morphologically diagnosable sibling species of the Diorhabda elongata species group are specialized feeders upon Tamarix. Four of these species, previously classified as D. elongata, have been released into the open field in the United States: D. carinulata, D. elongata, D. carinata, and D. sublineata. Diorhabda carinulata and D. elongata are confirmed as established in the U.S. Diorhabda meridionalis has yet to be cultured for study in the U.S. a) As distinct species, members of the D. elongata group are strongly reproductively isolated and would interbreed rarely, if at all, in the open field whether in the Palearctic or North America. b) Laboratory or cage produced interspecific hybrids of these species probably all experience varying degrees of hybrid breakdown which would probably lead to poor persistence in the open field and render them unsuitable in tamarisk biological control, especially in comparison to parental pure lines. However, the degree of breakdown in backcross hybrids may be lower than in hybrid/hybrid crosses and has yet to be studied.

2) Each of the five species of the D. elongata group has unique biogeographic characteristics in the Old World, even those species occurring in partial sympatry. These differences are probably related to innate biological characteristics of these species that would probably also lead to unique distributional patterns in the New World. a) Each member of the D. elongata group is probably uniquely suited to different ecoregions of the North American tamarisk invasion. b) Diorhabda carinulata, D. carinata, D. sublineata and D. meridionalis all appear to be better suited to differing desert and grassland ecoregions of southwestern North America than is D. elongata, primarily a species of the maritime Mediterranean biome. Efforts are needed to establish both D. carinata and D.

128 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS sublineata at various sites in west Texas in order to compare their efficacy in controlling tamarisk with that of D. elongata.

3) Previous host range testing demonstrates that D. carinata and D. sublineata (both recently introduced into the open field in Texas) are just as safe in terms of risks to non-target plants of Tamarix aphylla and Frankenia species as are the species D. carinulata and D. elongata.

4) Southern climatypes of D. carinata and D. carinulata from southwest Asia may be better pre-adapted to latitudes from ca. 30–34°N in North America than current northern climatypes of these species in the U.S.

5) Based upon its native distribution, Diorhabda meridionalis warrants host-range testing as a potential tamarisk biological control agent that may be the best suited tamarisk beetle to subtropical maritime desert ecoregions of North America.

The morphological and distributional evidence for reproductive isolation between the five species of the D. elongata group, including the lack of intermediate hybrid morphologies in nature, has been thoroughly discussed previously in this revision under the species accounts. Our Habitat Suitability Index (HSI) model for the D. elongata group in North America depicts how each of the five tamarisk beetles appear to be uniquely suited to different tamarisk infested ecoregions of western North America (Map 13). The HSI model estimates D. elongata as best suited to the Mediterranean biome of northern California, and four other Diorhabda species as better suited than D. elongata to grassland and desert biomes across the southwest (Map 13). The wide range of geographic areas occupied by the D. elongata species group provides great potential not only for interspecific variation in biogeographical traits, but also for intraspecific variability in critical traits such as climatic adaptation and Tamarix host preferences (Maps 1 and 8). Intraspecific genetic variants may take the form of climatypes and Tamarix host ecotypes. Climatypes can differ in synchrony of seasonal changes in physiology and behavior with environmental cues such as daylength and temperature. Geographic (allopatric and parapatric) climatypes are known among several insect species (Leather et al. 1993, Shapiro 1995, Singer et al. 1995) and sympatric climatypes, in the form of voltinism ecotypes, among at least one species (McLeod et al. 1979, Coates et al. 2004). Bean and Keller (in prep) found evidence for climatypes in intraspecific populations of Diorhabda carinulata (as D. e. deserticola) from Fukang and Turpan, China which differ in the critical photoperiod for diapause induction. In chrysomelids, Hsiao (1978) has reported intraspecific geographic host ecotypes and Ikonen et al. (2003) has reported sympatric host ecotypes. The value of intraspecific genetic variability found in differing “strains” of biological control agents has been challenged, mainly regarding biological control agents of (Clarke and Walter 1995), and more research is needed. But, selection of ecotypes can be critical in successful biological control, because not all populations may carry the genetic variation for desired traits (Zwölfer and Preiss 1983, Zwölfer and Harris 1984, Luck et al. 1995, Goolsby et al. 2006). Established climatypes may gradually adapt to wider areas within the biogeographic range of the species as appears to be happening with the northern climatype of D. carinulata which may have potential to establish further south in eastern New Mexico. Tamarisk beetle climatypes that are mismatched to the area of introduction may establish more slowly, or not at all. A climatypic mismatch may have contributed to delayed establishment of D. elongata introduced from the warm climates of coastal Crete into northern California. For tamarisk beetle species with native distributions over large latitudinal/climatic gradients, such as D. carinata and D. carinulata, collecting, host range testing and introducing two or three putative climatypes from throughout the latitudinal/climatic range of each species might increase or speed establishment success at corresponding tamarisk habitats in North America. A northern climatype of D. carinulata from northwest China and Kazakhstan has defoliated large acreages of tamarisk and is showing potential for substantial tamarisk suppression in the continental temperate cold deserts of the western U.S. Introduction of a southern climatype of northern tamarisk beetles

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 129 from Iran and Pakistan (Map 6) might facilitate more rapid and complete range expansion into its potential range in the southern half of the Colorado Plateau Shrublands, western portions of the Mojave Desert, the Trans-Pecos Chihuahuan Desert, and southwestern Western Short Grasslands (Map 13). Diorhabda elongata from Greece shows excellent promise in defoliating T. chinensis × T. canariensis/T. gallica in the Western Short Grasslands of west Texas and T. parviflora in the California Interior Chaparral and Woodlands of California. The biomic profile of D. elongata best matches T. parviflora and T. gallica among invasive North American tamarisks. Mediterranean tamarisk beetles may be best suited to maritime temperate warm Mediterranean ecoregions in central and northern California where T. parviflora is primarily invasive (Maps 7, 9). Southern climatypes of D. carinulata and D. carinata and the species D. meridionalis and D. sublineata are probably best suited to desert and grassland ecoregions of the southwestern U.S. and could probably replace D. elongata where it becomes established in these ecoregions. The biomic profiles of D. carinulata and D. carinata most closely match that of the target weed T. ramosissima. The northern Qarshi climatype of D. carinata is probably most suitable to warm temperate grasslands of western Kansas and eastern Colorado. Introduction of a southern climatype of D. carinata from Iraq, eastern Iran or central Pakistan (Map 6) might facilitate more rapid and complete range expansion of this species from 31–37°N in deserts and grasslands such as southern portion of the Western Short Grasslands and Central and Southern Mixed Grasslands, the southern portion of the Colorado Plateau Shrublands, the Trans-Pecos Chihuahuan Desert, and northern portions of the Mojave Desert, (Map 13). Diorhabda sublineata is probably the best suited tamarisk beetle for several Mediterranean and maritime subtropical desert areas of various ecoregions, including the California Coastal Sage and Chaparral, southern portions of the Chihuahuan, Mojave and Sonoran deserts, and Tamaulipan Mezquital (Map 13). Diorhabda meridionalis (not yet cultured) from southwest Asia is a maritime subtropical desert species that may be uniquely adapted to parts of the Sonoran Desert and Tamaulipan Mezquital (Map 13). If D. sublineata does not establish well in portions of the Sonoran deserts in extreme southern California and the Tamaulipan Mezquital in south Texas, investigation into the possibility of introducing D. meridionalis into these areas might be warranted. Firmly establishing D. carinata and D. sublineata in the field, and possibly acquiring D. meridionalis and southern climatypes of D. carinulata and D. carinata for testing and introduction, could expand and speed the success of tamarisk biological control across the southwestern U.S. Interspecific differences in biogeographic preferences of these species are probably not solely the result of variation in synchronization of diapause induction with local photoperiodic and climatic conditions (climatypic traits which can also widely vary intraspecifically), but are also likely to be related to differences in adaptation to regional variation in a number of climatic variables, including temperature, relative humidity, and precipitation. Introduction of tamarisk beetle species at North American locations to which they are biogeographically mismatched may result in failed or poor establishment with population crashes or eventual extinction under climatic extremes to which the species are ill-adapted. The poor establishment of D. elongata in the northwestern Chihuahuan Desert near Artesia, New Mexico, may be an example of a biogeographic mismatch, as the Mediterranean tamarisk beetle is not known from deserts in its native habitat. Species distribution models (SDMs) can be useful tools in estimating potential ranges of biological control agents of weeds in non-native areas (McFayden 1991). SDMs based on the presence-only locality data from this revision and incorporating a variety of global climatic geographic information system data (e.g., Elith et al. 2006) could be used to estimate potential differences in North American distribution of these tamarisk beetles. Studies in field cages are in progress to evaluate diapause characteristics, voltinism, and overwintering survival across several latitudinal gradients in the western U.S. for populations of D. elongata (from Posidi Beach and Sfakaki, Greece), D. carinulata (from Fukang and Turpan, China), D. sublineata (from near Sfax, Tunisia), and D. carinata (from near Qarshi, Uzbekistan) (Peter Dalin and Tom Dudley, University of California, Santa Barbara, CA, pers. comm.). Results from these field studies should be compared to those of species distribution models. At present, laboratory and field studies lack access to populations of D. meridionalis and putative southern climatypes of D. carinulata and D. carinata.

130 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Host range safety testing was conducted for each differing tamarisk beetle population/species prior to release (DeLoach et al. 2003b; Lewis et al. 2003a; Milbrath and DeLoach 2006a, 2006b; Milbrath et al. 2007; Herr et al. 2006, in prep.). All four tested tamarisk beetle species (D. carinulata, D. elongata, D. carinata and D. sublineata) represent very low risk of damaging North American Frankenia spp. (Lewis et al. 2003a; Dudley and Kazmer 2005; Milbrath and DeLoach 2006a; Dudley et al. 2006; Herr et al. 2006, in prep.) and a moderate risk of damaging T. aphylla (DeLoach et al. 2003b, Milbrath and DeLoach 2006b). Several Frankenia species are indigenous throughout the Palearctic (Jäger 1992) and overlap in distribution with the D. elongata group where they can also be found together in the same habitat as Tamarix (Kassas and Imam 1954). However, no Frankenia spp. are recorded hosts of Diorhabda in the Palearctic (Table 1). Surveys of Frankenia spp. and Tamarix spp. in Tunisia by our cooperators R. Sobhian and A. Kirk (USDA-ARS European Biological Control Laboratory [EBCL], Montferrier-sur-Lez, France) revealed D. sublineata only on Tamarix (DeLoach et al. 2003b). Frankenia appears only to serve as a factitious, artificial laboratory host, but not a natural host for the four studied species of tamarisk beetles. High populations of tamarisk beetles in the early stages of biological control may pose a transitory risk in damaging leaves of Frankenia growing in proximity to tamarisk (see Dudley and Kazmer 2005). The ornamental T. aphylla is generally less preferred for oviposition over other Tamarix in field cage preference studies of the four species of tamarisk beetles (DeLoach et al. 2003b; Milbrath and DeLoach 2006a, 2006b). Diorhabda elongata clearly prefers T. ramosissima/T. chinensis over T. aphylla for oviposition in the open field at Big Spring, Texas (Herr et al. 2006, Moran et al. in press). However, T. aphylla is accepted equally well as T. ramosissima × T. chinensis in no-choice tests of three species of tamarisk beetles (Milbrath and DeLoach 2006b). Tamarix aphylla is uncommonly reported as a native host of any tamarisk beetle (Table 1), and we expect that any damage to T. aphylla from the four tested tamarisk beetles will be uncommon compared to damage on other tested invasive deciduous Tamarix spp. The recent recognition of T. aphylla as an invasive tree in part of the southwestern U.S. (Walker et al. 2006) should weigh against concerns over possible damage to T. aphylla from tamarisk beetles. Only the southern extremes of the indigenous ranges of D. carinulata and D. elongata overlap with the distribution of T. aphylla in the Palearctic realm. We expect that the ranges of D. elongata and D. carinulata would also rarely come into contact with T. aphylla in North America. The benefits of tamarisk biological control with tamarisk beetles outweigh the above described risks to nontarget plants and other environmental risks (DeLoach 1990, DeLoach et al. 2000, Dudley et al. 2000). Additional field studies from both native and introduced habitats should provide better understanding of interspecific differences in Tamarix host preferences among the D. elongata group. Studies across the range of each species of tamarisk beetle also may reveal intraspecific variations in Tamarix preferences. Diorhabda carinulata is best known in terms of field host preferences. It reportedly prefers T. ramosissima over much of its northern and eastern range in Asia, but the Tamarix spp. preferred in its southern and western range are unknown. Diorhabda carinulata appears to have a fairly broad host range within the genus Tamarix, as evidenced by its feeding on a novel host, T. parviflora, in Nevada, but T. ramosissima is attacked more than T. parviflora (Dudley et al. 2006). Diorhabda elongata is predictably beginning to defoliate one of its natural hosts, T. parviflora, in California. Diorhabda elongata attacks T. smyrnensis, a close relative of T. ramosissima, and T. gallica, which may account for its successful adaptation to T. chinensis × T. canariensis/ T. gallica in west Texas. Diorhabda carinata is common on T. arceuthoides and T. hispida in Tajikistan and it occurs on T. ramosissima among several other Tamarix spp. in central Asia, but its field host preferences are unknown. Hosts of D. sublineata include T. gallica, which comprises part of the T. canariensis/T. gallica complex that commonly hybridizes with T. ramosissima/T. chinensis in Texas. Very little is known of the Tamarix hosts of D. meridionalis. Multiple agents targeting a weed can improve the efficacy of biological control, especially where agents of differing habitat/climatic adaptations are needed and attack of multiple plant parts (e.g., roots and foliage) is achieved. These benefits must be weighed against additional risks to non-target plants (in this case Frankenia spp. and T. aphylla) and possibly deleterious competitive interactions of the agents (Denoth et al. 2002). Potentially counter-productive indirect competition between multiple weed biological control agents

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 131 has been observed in several programs, such as between two weevil species used in biological control of the exotic biennial weed musk thistle, Carduus nutans Linnaeus (Milbrath and Nechols 2004a, 2004b). On the other hand, many examples are known of multiple agents being successfully used together in biological control over the long term, especially against perennial weeds (Blossey and Hunt-Joshi 2003). Several species of root-mining Aphthona spp. flea beetles (Chrysomelidae: Galerucinae) with differing habitat preferences are being successfully used in biological control of exotic leafy spurge, Euphorbia esula Linnaeus, in North America (Gassmann et al. 1996, Nowierski et al. 2002). The syntopic closely related chrysomelids (Linnaeus) and G. pusilla (Duftschmidt) appear to be equal in their competitiveness and are fully complementary in their biological control of purple loosestrife, Linnaeus, in North America (Blossey 1995). Potential competitive interactions among species of the D. elongata group are yet to be studied, but would probably be limited to species occupying similar biogeographic ranges. Biogeographic relationships found among species of tamarisk beetles in their native range (Table 8) would probably also apply to their ranges in North America. The sympatric and syntopic species pairs of D. carinata/D. carinulata and D. carinata/D. meridionalis might also be syntopic over partially sympatric ranges in North America. In differing areas of the Old World, one member of these partially sympatric species pairs appears to be generally more abundant than the other. Near Ashgabat, Turkmenistan, D. carinata successfully defoliates tamarisk at Dry Sport Lake in the presence of smaller syntopic populations of D. carinulata. Likewise, near Shelek, Kazakhstan, D. carinulata defoliates tamarisk in the presence of smaller syntopic populations of D. carinata. In North America, the efficacy of tamarisk biological control would probably not be reduced where both species of these sympatric/ syntopic species pairs are present. Diorhabda carinulata and D. sublineata are allopatric in their native range and probably would also be allopatric in North America. Species pairs that are parapatric or marginally sympatric in the Old World (Table 8) might compete where their ranges contact in North America, but areas of range contact are likely to be small as a result of interspecific differences in biogeographic traits. Marginally sympatric species pairs in the Palearctic with potential for range contact in North America include D. elongata/D. carinulata, D. elongata/D. sublineata, D. elongata/D. carinata, and D. sublineata/D. carinata.In the small areas of range contact between these potentially competitive species, one species is likely to predominate with probably no reduction in efficacy for tamarisk biological control. This appears to be the case in the Old World where one of these species is generally dominant in the area of marginal sympatry with another species (Map 1, Table 8). Studies are in progress to evaluate potential competitive interactions between D. elongata and D. carinulata in field cages in New Mexico (D. C. Thompson, pers. comm.). Several species of tamarisk beetles hybridize readily in the laboratory, producing hybrids with varying levels of reduced egg viability in their progeny. However, the lack of detected morphological hybrid forms in the Old World provides no evidence that different tamarisk beetles hybridize in nature. Differing mate recognition systems appear to prevent or severely reduce field hybridization of tamarisk beetle species. Uncommon instances of interspecific hybridization with the production of fertile hybrids might occur among different tamarisk beetles in areas of sympatry. Laboratory hybridization of D. carinulata and D. carinata leads to severely reduced hybrid F1 and F2 egg viabilities and high mortality in copulo for matings of D. carinata males and D. carinulata females. However, these two species inhabit the same tamarisk trees in parts of central Asia with no apparent interference from hybridization in defoliating tamarisk, either for D. carinata in Ashgabat, Turkmenistan, or for D. carinulata in Chilik, Kazakhstan. Consequently, these two species should be able to similarly coexist without interference in North America and the same probably applies to all members of the D. elongata group. The production of uncommon fertile hybrids in nature is seen in some animal species (Mallet 2005), but Helbig et al. (2000) point out the lack of any known cases of complete breakdown in reproductive isolation between any animal species. Recently, evidence of hybrid speciation, a potential means of adaptive radiation when populations invade new environments (Seehausen 2004), has been found among closely related insects (Salazar et al. 2005, Schwarz et al. 2005, Mavárez et al. 2006), but the frequency and occurrence of hybrid speciation among different insects is poorly known.

132 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Differing interspecific biogeographic preferences may be a critical factor in determining where each type of tamarisk beetle can establish in North America. Synchronization of diapause to daylength and temperature in response to variations in latitude and altitude has been a major focus in exploring factors affecting establishment success of Diorhabda in North America. However, biogeographic adaptations to variation in bioclimatic conditions across biomes at a given latitude and altitude, such as between desert riparian habitat and Mediterranean riparian habitat, are probably just as important in influencing establishment success. Nowierski et al. (2002) found that prior knowledge of the habitat preferences of Aphthona spp. flea beetles could have prevented delays in successful biological control of leafy spurge that resulted from trying to establish certain Aphthona species in non-suitable habitats in North America. Optimal matching of different tamarisk beetles to North American locations should involve both biogeographic matching at the species level and climatypic matching within each species. Within the biogeographic range of each species of tamarisk beetle, intraspecific climatypes are probably adapted to differing daylength/climatic regimes. Introduction of southern climatypes of D. carinulata and D. carinata from 30–34°N in southwest Asia (Map 6) might speed establishment of these species at corresponding latitudes in biogeographically suitable areas of the southwestern U.S (Map 13). At some sites in southwestern U.S., potentially several species of Diorhabda might be able to establish (Map 13), but probably one species is the best biogeographically suited and would establish more vigorously, reach greater population levels over time, and produce more rapid defoliation of tamarisk. For example, the maritime Mediterranean species D. elongata is established at several sites in the Trans-Pecos Chihuahuan Desert in west Texas, but our HSI models predict that the desert/grassland species D. carinata or D. carinulata would perform better at these sites. These HSI models are only a first rough approximation at predicting which species may perform best at any given location and our detailed distribution data should now be used to make further ecogeographic analyses using species distribution models with climatic data. Such models could both speed the process and enhance the prospects of matching appropriate tamarisk beetle species/climatype combinations to specific regions of the North American tamarisk invasion. Biological control has permanently reduced populations of many target weeds below the economic thresholds justifying use of expensive chemical and mechanical controls (DeLoach 1997, Gould and DeLoach 2002). Biological control ideally should be pursued as the cornerstone of modern integrated pest management programs (IPM) with the aim of reducing the need for chemical controls (O’Neil et al. 2003), and not considered primarily as only a follow up treatment to other controls. Integration of biological and chemical controls is appropriate in some weed management situations, but chemical controls can interfere with successful biological control (Ainsworth 2003) while providing less effective control in the long run than would biological control alone (e.g., Larson et al. 2007). Chemical control of tamarisk is extensively and aggressively being pursued in the U.S., especially in Texas and New Mexico. In California, Nevada, Utah, Wyoming, Colorado and Texas, tamarisk biological control with D. carinulata and D. elongata is showing good potential for gradually (over several years) producing sustained reductions in tamarisk stands to levels below the need for chemical control and with no harm to native vegetation. Additional efforts are needed to compare in the field the efficacy of D. carinata and D. sublineata against D. elongata as tamarisk biological control agents in southwestern deserts. More widespread demonstration of the effectiveness, ecological benefits, and low cost of tamarisk biological control in the southwestern U.S. is essential in order for biological control to be recognized as a primary management alternative to chemical control of tamarisk. The geographic extent, rapidity, and effectiveness of tamarisk biological control across the Southwest can potentially benefit from wider utilization of the biogeographic diversity and uniqueness of all five species of tamarisk beetles.

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 133 Opportunities for Further Research

Throughout this monograph, we refer to several studies in progress and suggest several areas for further study which we summarize below:

Category Research Task Researchers with Potential Interest Biological Compare efficacy (ease of establishment and rapidity of spread and Mark Muegge, Allen Control control of tamarisk) of D. elongata, D. carinata, and D. sublineata Knutson, Dave Thompson, in deserts and grasslands of southwestern North America Tom Dudley Taxonomy Search for external diagnostic characters in D. elongata group Dave Thompson, Jessica Perez, James Tracy Analyze external morphometrics of D. elongata group Joaquin Sanabria, James Tracy Characterize three dimensional morphology of inflated endophalli obtained from freshly killed mating pairs in the D. elongata group Search for taxonomically diagnostic characters at both species and genus level in male endophallic sclerites, female vaginal palpi, and interanal sternite VIII among Diorhabda and related genera Hybridization Publish crossing studies of D. elongata group (Thompson et al. in Dave Thompson, Dan Bean, prep.), and cross additional species, especially D. sublineata and D. Julie Keller, James Tracy, carinata Jack DeLoach Publish studies of mitochondrial and nuclear DNA in D. elongata, Dave Kazmer D. carinata, D. sublineata, and, D. carinulata. Use genetic and morphological techniques to look for potential hybrids in field population where species potentially interface, such as between D. sublineata and D. elongata in western Italy Biogeography Publish bioclimatic models to species distributions. James Tracy Better determine species distribution of D. elongata group in potential interspecific interface areas such as in western Italy and central Turkey Biology Publish data on critical photoperiod for D. elongata, D. carinata Dan Bean and Ray Carruthers and D. sublineata (Bean and Keller in prep.) (in prep.) Publish field data comparing host suitability (Dalin et al. in press) Peter Dalin, Tom Dudley, and site adaptability between D. carinulata, D. elongata, and D. Dave Thompson carinata Confirm pheromones for D. elongata, D. carinata, and D. Allard Cossé, Tom Dudley sublineata with field testing Publish field data and models for seasonal dispersal and defoliation Jack DeLoach and Joaquin by D. elongata Sanabria Determine native species of Tamarix serving as hosts for D. meridionalis Perform host range testing for D. meridionalis Obtain southern climatypes of D. carinulata from around 31°N in Dan Bean Pakistan or Iran and compare critical daylength with northern populations

134 · Zootaxa 2101 © 2009 Magnolia Press TRACY & ROBBINS Acknowledgements

We are first grateful to our friend and mentor C. Jack DeLoach (USDA-ARS, Temple, Texas), both for his insightful review of this manuscript and his prodigious and inspiring effort in initiating the tamarisk biological control program with Diorhabda in the United States and his continued efforts in research, implementation and monitoring of tamarisk biological control with associated ecosystem effects in Texas. We are grateful to Kuniko Arakawa (Moriya–shi, Japan) for excellent illustrations of the genitalia in males (Figs. 14–18) and females (Figs. 34–38) of the five species of tamarisk beetles. We especially thank Ed Riley (Texas A & M University, College Station, TX) and Alexander Konstantinov for their ready encouragement and technical advice throughout this revision. We are also grateful to Pierre Jolivet (Paris, France) for helpful suggestions and providing copies of color plates from Brullé (1832). We thank Joaquin Sanabria (Texas Agricultural Experiment Station, Temple, TX) for assistance with statistical analyses. We thank Nell Messer, Travis Sawin, Chris Kroll, Chris Elias, Emily Martinez, and Erin Albright for assistance in entry of specimen label data, researching geocoordinates of collection locations, and specimen labeling. We are grateful to Jeri Palmer and Chris Puetsch for assistance with georeferencing Old World tamarisk distribution maps and researching geocoordinates of tamarisk locations. We appreciate testing of early versions of the key to males by Lindsey Milbrath (USDA-ARS, Ithaca, NY) and Ed Riley. We express our sincere appreciation to our colleagues who collected the original material for live cultures of the beetles – Rouhollah Sobhian and Alan Kirk (USDA- ARS EBCL, Montferrier-sur-Lez, France, retired), Javid Kashefi (USDA-ARS EBCL, Thessaloniki, Greece), Massimo Cristofaro (ENEA C.R. Casaccia, Rome, Italy) and Ray Carruthers (USDA-ARS, Albany, CA). We thank Dan Bean (Colorado State Department of Agriculture, Grand Junction, CO), Tammy Wang (USDA- ARS, Albany, California), John Herr (USDA-ARS, Albany, CA), Dave Thompson (New Mexico State University, Las Cruces, NM), and Debra Eberts (USDI Bureau of Reclamation, Denver, CO) for supplying us with live cultures of Diorhabda species. We are also grateful to Dan Bean (Colorado State Department of Agriculture, Grand Junction, CO), Patrick Moran (USDA-ARS, Weslaco, TX), and Ray Carruthers (USDA- ARS, Albany, TX) for organizing symposiums where this and other research on tamarisk beetles could be presented and discussed. We thank Igor Lopatin, Alexander Konstantinov, Steven Lingafelter, Shawn Clark, and Richard White for earlier taxonomic determinations of specimens from our collections. We appreciate John Gaskin (USDA-ARS, Sidney, MT) for DNA analysis and identification of Tamarix specimens. We thank G.M. Thomas (Berkeley, CA), T. Poprawksi (USDA-ARS, Weslaco, TX, deceased) and J. Siegel (USDA- ARS, Parlier, CA) for identifications of pathogens of Diorhabda. We are indebted to the following curators of the various museums that provided us loans of specimens used in this revision: Ben Brugge (ZMAN), Mauro Daccordi (MRSN), Roy Danielsson (MZLU), Ariel-Leib-Leonid Friedman (TAU), Johannes Frisch (ZMHB), Jiri Hajek (NMPC), Alexander Konstantinov (USNM), Boris Korotyaev (ZIN), Pol Limbourg (IRSNB), Carolina Martín (MNMS), Otto Merkl (HNHM), Nailya Mirzoeva (AMA), M. Ashraf Poswal (CABP), Renato Regalin (MSNM), Hans Riefenstahl (ZMUH), Sharon Shute (BMNH), Eva Sprecher (NHMB), Hans Silfverberg (MZHF), Dave Thompson (NMSU), Bert Viklund (NHRS), Xingke Yang (IZAS), and Lothar Zerche (DEI). We appreciate the following individuals who allowed us to examine specimens from their personal collections: Edward G. Riley (EGRC), İrfan Aslan (IAET), Michel Bergeal (MBPF), and Ron Beenen (RBCN). We thank the following individuals who provided Old World specimens for our collection (GSWRL) used in this revision: Paul Boldt (USDA/ARS, Temple, TX, retired), Bao Ping Li (Nanjing Agricultural University, Nanjing, China), John Gaskin, Roman Jashenko and Ivan Mityaev (Tethys Scientific Society/Kazakhstan Academy of Science, Almaty, Kazakhstan), Javid Kashefi, Alan Kirk, Lev Medvedev (Russian Academy of Sciences, Moskva, Russia), Svetlana Myartseva (Ashgabat, Turkmenistan), Ilya Osipov (Feasterville, PA), Sassi (Casterlmarte, Italy), Rouhollah Sobhian, Jose Vela (Malaga, Spain), and Andrzej Warchalowski (Instytut Zoologicny UWr.,Wroclaw, Poland). We thank the following people who provided specimens from United States field sites where beetles were introduced and established: Tom Dudley (Univ. CA, Santa Barbara), Dave Kazmer (USDA-ARS, Sidney, MT), Fred Nibling (US Bureau of Reclamation [BR], Denver, CO), Denise Hosler (USBR, Denver, CO), Debra Eberts (USBR, Denver, CO),

DIORHABDA ELONGATA SPECIES GROUP Zootaxa 2101 © 2009 Magnolia Press · 135 Jerry Michels, Vanessa Carney, Erin Jones (Texas AgriLIFE Research, Amarillo, TX), and Charles Randal (USDA-APHIS, Olney, TX). We appreciate the following individuals who provided unpublished locality data for Diorhabda used in our revision: Serge Doguet (Fontenay–sous–Bois, France), Alan Kirk, and Bao Ping (Alashan Range Extension Station, Nei Mongol Zizhiqu, China). We thank Zhang Peng-yun (Lanzhou University, Lanzhou, Gansu Sheng, China) for unpublished locality data for T. austromongolica. We appreciate personal communications of unpublished data on tamarisk beetles regarding life history by Lindsey Milbrath, interspecific hybridization by Beth Peterson (New Mexico State University, Las Cruces, NM), Dave Thompson, Dan Bean and Julie Keller, DNA comparisons by Dave Kazmer, diapause induction by Dan Bean, putative pheromones by Robert Bartelt (USDA-ARS, Peoria, IL), and field ecology and behavior by Ray Carruthers (USDA-ARS, Albany, CA), Tom Dudley (University of California, Santa Barbara, CA), Allen Knutson (The Texas AgriLIFE Extension Service, Dallas, TX), Mark Muegge (The Texas AgriLIFE Extension Service, Fort Stockton, TX), Dan Bean (Colorado Department of Agriculture, Grand Junction, CO), Mark Donet (USDA-NRCS, Alpine, TX), Andrew Berezin (Sul Ross State University, Alpine, TX), and Charles Randal. We thank several who assisted by translating material into English from various languages including, Bao Ping Li, Hongyin Chen, Benjamin Sheng, Svetlana Myartseva, Roman Jashenko, Irina Kashefi, Lothar Zerche, Mauro DiLuzio, Pierre Jolivet, Alexander Konstantinov, Martin Volk, and Anna Mkhitaryan. We are grateful to Ed Riley, Alexander Konstantinov, Shawn Clark, Mauro DiLuzio, and Chadwick Rittenhouse for reviewing the manuscript. We especially thank Juliett Meleshko (Byelorussian State University, Minsk, Belarus) and Roman Jashenko for preparation of the Russian language abstract. We also thank anonymous reviewers for their valuable comments on the manuscript. This research was funded in part by a USDA-CSREES-IFAFS grant #00-52103-9647 and Texas State Soil and Water Conservation Board Project 03-11.

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